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g° t » NEWSLETTER 

Number 50 16.11.2012 



Scientific Editor: Bertil Nordenstam 

Technical Editor: Gunnel Wirenius Nohlin 

Published and distributed by The Swedish Museum of Natural History, 

Department of Phanerogamic Botany, 

P.O. Box 50007, SE-104 05 Stockholm, Sweden 

ISSN 0284-8422 



Contents 



Editorial: On the magic number 50 

Funk, V. A. & P. Pelser: The Compositae Newsletter reaches its 

50th volume and Bertil Nordenstam steps down as Editor after 25 years 4 

Funk, V. A. & L. Brouillet: News from The International Compositae 
Alliance 2012 Meeting 9 

Brouillet, L. & V. A. Funk: The International Compositae Alliance 

(TIC A) Montreal, Canada 15-19 July 20 1 2 (Abstracts and Posters) 1 4 

Anderberg, A. A.: On the identity of Pluchea incisa Elmer, with notes 

on the genera Blumea and Pluchea (Asteraceae-Inuleae) 36 

Anderberg, A. A.: Ray-florets in Chiliadenus (Asteraceae-Inuleae), 
discovered. An epigenetic phenomenon? 40 

Anderberg, A. A. & J. I. Ohlson: The genus Cavea, an addition to the 

tribe Gymnarrheneae (Asteraceae-Gymnarrhenoideae) 46 

Nordenstam, B. & P. B. Pelser: Caputia, a new genus to accommodate 

four succulent South African Senecioneae (Compositae) species 56 

Nordenstam, B.: Crassothonna B. Nord., a new African genus of 

succulent Compositae-Senecioneae 70 

Mukherjee, S. K. & B. Nordenstam: Diversity of trichomes from mature 
cypselar surface of some taxa from the basal tribes of Compositae 78 

New taxa and combinations published in this issue 126 



Comp.Newsl. 50,2012 



Editorial 
On the magic number 50 

In nuclear physics the number 50 is one of seven so-called magic numbers, and they 
are involved with nuclear isotope stability. In the plant world there is no such thing 
as magic numbers connected with stability, although the frequent occurrences of 
somewhat magic Fibonacci numbers are noted in the Compositae, e.g., in number 
of phyllaries, ray-florets, disc-florets etc. Sunflowers and artichokes are well 
known examples. These numbers are part of the Fibonacci series 0, 1, 1, 2, 3, 5, 
8, 13, 21, 34, 55 etc., and there is no room for 50 there. Also, although strived for, 
stability is not as obvious in taxonomy as maybe in nuclear physics. 

Compositae Newsletter No. 50 

To me, at this moment, the number 50 nevertheless has some magic. This is the 
final issue of the Compositae Newsletter, at least under my editorship, and it bears 
the number 50. 

50 years of research on Compositae 

My own research on the family has been ongoing for >50 years. In 1962 I went 
to South Africa for a two-year period of field and herbarium work, as the first 
non-South African Smuts Memorial Fellow. Although I have visited more than 75 
countries in all continents (except Antarctica), Southern Africa remained my main 
hunting grounds for plants, with another half-year stay in 1 974 and several shorter 
later visits - in all about three years. 

50 new Compositae genera 

By magic coincidence I have now published 50 names of new genera of 
Compositae. Some have gone into synonymy, e.g. due to refined knowledge on 
phylogeny especially through molecular studies, but most of them are still valid 
and in current use. Here is the list of the genera in alphabetical order. 

l.Acrisione B. Nord.- Bot. Jahrb. Syst. 107(1-4): 582 (1985) 

2. Adenanthellum B. Nord. - Bot. Not. 132(2): 160 (1979) 

3. Adenanthemum B. Nord. - Bot. Not. 129(2): 157 (1976) 

4. Adenoglossa B. Nord. - Bot. Not. 129(2): 137 (1976) 



2 Comp.Newsl. 50,2012 

5. Aequatorium B. Nord. - Opera Bot. 44: 59 (1978) 

6. Anderbergia B. Nord. -Ann. Naturhist. Mus. Wien 98B(Suppl.): 407 (1996) 

7. Antillanthus B. Nord. - Compositae Newslett. 44: 51 (2006) 

8. Canariothamnus B. Nord. - Compositae Newslett. 44: 26 (2006) 

9. Capelio B. Nord. - Compositae Newslett. 38: 72 (2002) 

10. Caputia B. Nord. & Pelser - Compositae Newslett. 50: (2012) 

11. Caucasalia B. Nord. -PI. Syst. Evol. 206(1-4): 22. (1997) 

12. Comptonanthus B. Nord. - J. S. African Bot. 30: 54 (1964) 

13. Crassothonna B. Nord. - Compositae Newslett. 50: (2012) 

14. Cymbopappus B. Nord. - Bot. Not. 129(2): 150 (1976) 

15. Dauresia B. Nord. & Pelser - Compositae Newslett. 42: 76 (2005) 

16. Dendrosenecio (Hauman ex. Hedberg) B. Nord. - Opera Bot. 44: 40 (1978) 

17. Dolichoglottis B. Nord. - Opera Bot. 44: 33 (1978) 

18. Elekmania B. Nord. - Compositae Newslett. 44: 66 (2006) 

19. Graphistylis B. Nord. - Opera Bot. 44: 56 (1978) 

20. Herreranthus B. Nord. - Compositae Newslett. 44: 62 (2006) 

21. Hilliardia B. Nord. - Opera Bot. 92: 147 (1987) 

22. Igmtrbia B. Nord. - Willdenowia 36( 1 ): 464 (2006) 

23. Imiloides B. Nord. - Compositae Newslett. 44: 44 (2006) 

24. Io B. Nord. - Compositae Newslett. 40: 47 (2003) 

25. Iocenes B. Nord. - Opera Bot. 44: 58 (1978) 

26. Iranecio B. Nord. -Fl. Iranica [Rechinger] 164: 53 (1989) 

27. Jacmaia B. Nord. - Opera Bot. 44: 64 (1978) 

28. Lamprocephalm B. Nord. - Bot. Not. 128(3): 323 (1976) 

29. Leonis B. Nord. - Compositae Newslett. 44: 55 (2006) 

30. Leucoptera B. Nord. - Bot. Not. 129(2): 140 (1976) 

3 1 . Lomanthus B. Nord. & Pelser - Compositae Newslett. 47: 34 (2009) 

32. Lordhowea B. Nord. - Opera Bot. 44: 38 (1978) 



Comp. Newsl. 50, 2012 3 

33. Lundinia B. Nord. - Compositae Newslett. 44: 64 (2006) 

34. Monoculus B. Nord. - Compositae Newslett. 44: 39 (2006) 

35. Nemosenecio (Kitam.) B. Nord. - Opera Bot. 44: 45 (1978) 

36. Nephrotheca B. Nord. & Kallersjo - Compositae Newslett. 44: 33 (2006) 

37. Nesampelos B. Nord. - Compositae Newslett. 45: 37 (2007) 

38. Norlindhia B. Nord. - Compositae Newslett. 44: 41 (2006) 

39. Notoniopsis B. Nord. - Opera Bot. 44: 69 (1978) 

40. Odontocline B. Nord. - Opera Bot. 44: 23 (1978) 

41. Oldfeltia B. Nord. & Lundin - Compositae Newslett. 38: 66 (2002) 

42. Oresbia Cron & B. Nord. - Novon 16(2): 216 (2006) 

43. Phaneroglossa B. Nord. - Opera Bot. 44: 66 (1978) 

44. Roodebergia B. Nord. -Acta Phytotax. Geobot. 53(2): 101 (2002) 

45. Scyphopappus B. Nord. - Bot. Not. 129(2): 147 (1976) 

46. Sinosenecio B. Nord. - Opera Bot. 44: 48 (1978) 

47. Stenops B. Nord. - Opera Bot. 44: 73 (1978) 

48. Urostemon B. Nord. - Opera Bot. 44: 31 (1978) 

49. Xyridopsis B. Nord. - Opera Bot. 44: 75 (1978) 

50. Zemisia B. Nord. - Compositae Newslett. 44: 72 (2006) 

About 20 of the genera are South African, and 16 are New World genera including 
the West Indies. Most of the new genera belong in the tribe Senecioneae, but 
some are members of other tribes such as the Calenduleae, Astereae, Anthemideae 
and Gnaphalieae. This is not the end, rather the end of a beginning. A few more 
genera of Senecioneae will have to be described, and the future of Compositae 
Newsletter is under discussion (see elsewhere in this issue). 

I wish to thank everybody involved in the Compositae Newsletter for support and 
contributions over the years - authors as well as subscribers and technical support 
first of all from Mrs. Gunnel Wirenius Nohlin and also from other staff in the 
Swedish Museum of Natural History. 

Bertil Nordenstam 



Comp. Newsl. 50, 2012 



The Compositae Newsletter reaches its 50 th 

volume and Bertil Nordenstam steps down as 

Editor after 25 years 



Vicki A. Funk 1 & Pieter Pelser 2 



'US National Herbarium, Department of Botany 

Smithsonian Institution, MRC 166 Washington, DC, USA 20013 

funkv@si.edu 



2 School of Biological Sciences, University of Canterbury, Private Bag 4800 
Christchurch 8140, New Zealand 

Professor Bertil Nordenstam (S) has been the editor of the Compositae Newsletter 
(Comp. Newsl.) for 25 years and he recently announced that he is stepping down 
after number 50 (this issue). 

The Compositae Newsletter began in 1975 and throughout its entire history it has 
always been free of charge. The scientific editing has been a "labor of love" by our 
colleagues and the technical editing, printing, and distribution has been supported 
by an institution or a donation. The first issue was a short announcement and the 
editors were Tod Stuessy (Ohio State University) and Robert M. King. It was 
organized by Stuessy and printed and mailed by the Smithsonian Institution. By 
the second issue Stuessy was the only editor and it was distributed by Ohio State 
University. Stuessy continued as editor until number 5 was published in 1980. 
For numbers 6-12 (1977 - 1982) it moved to Kew and Charles Jeffrey served 
as Editor. Numbers 4-12 received financial support from Mr. Sven Koeltz and 
numbers 9-12 were jointly sponsored by the Bentham-Moxon Trust at Kew. One 
issue, Number 13, was edited by Jette Baagoe, University of Copenhagen, and 
printed by the University of Stockholm out of the office of the Nordic Journal of 
Botany. For more information on the early history of the Compositae Newsletter 
see Nordenstam (1988). The first 13 numbers began as a true newsletter with 
information on recent publications, requests for samples, and announcements. 
Over the years it gradually added articles of general use such as lists of types, 
verification of names, and lists of literature pertaining to the Compositae. 

For six years no numbers of the Compositae Newsletter were published. Then, 
like a Phoenix rising from the ashes, Professor Bertil Nordenstam (S) brought it 
back to life in a new and improved form. It still included announcements and lists 



Comp. Newsl. 50, 2012 



of types but it also began to publish journal articles and comprehensive works. 
Some examples include Jansen et al.'s chloroplast DNA paper (1988), Bremer's 
discussion on corolla types (1988), Hind & Jeffrey's work on the Compositae 
of HBK (2001), Herrera & Ventosa's Cuban Asteraceae (2005), the Asteraceae 
of Chacoan Plain (Freire et al. 2005), and the molecular work on Dipterocome 
(Anderberg et al. 2007). It now has a well developed following and people 
regularly submit articles of interest. Other changes include its association with The 
International Compositae Alliance (TICA) and its move into the digital age: all 
issues are available free of charge through the Biodiversity Heritage Library web 
site: http://www.biodiversitylibrary.org/bibliography/1256 1 . This has increased 
its visibility and resulted in additional citations and submissions. All these things 
have been accomplished because of the efforts of Bertil Nordenstam and the 
technical editor Gunnel Wirenius Nohlin (starting with number 24, 1994). For 25 
years the journal has been supported by the Department of Phanerogamic Botany, 
Swedish Museum of Natural History, Stockholm, Sweden (S). Since 1988 they 
have provided a technical editor, and printed and mailed the journal to over 600 
scientists and libraries around the world, free of charge, but from 201 3 the museum 
financial support for technical editing and publishing cannot be maintained. Our 
colleagues at S have been more than generous for many years and we thank them 
very much. 

The Compositae Newsletter has published papers on the taxonomy of many 
Asteraceae taxa and one of these is Senecioneae, a tribe that is close to the heart 
of Bertil Nordenstam. His contributions to the Senecioneae date back to the 
start of his career (e.g., Nordenstam 1961) and are continuing until this very 
day. His papers often reported discoveries resulting from his extensive field work 
(Fig. 1). In this issue, two new Senecioneae genera are described: Caputia B. 
Nord. & Pelser and Crassothonna B. Nord., and we are informed that some more 
will follow. The Senecioneae publications in Compositae Newsletter are among 
the key literature on the tribe and include the publication of new genera (e.g., 
Nordenstam et al. 1997b, 2009; Nordenstam & Pelser 2005) and species (e.g., 
Nordenstam et al. 2002; Beltran & Baldeon 2009; Noroozi et al. 2010), new 
combinations (e.g., Nordenstam 1996, 1997, 1999; Beltran 1999; Pelser et al. 
2006), discussions of nomenclature and typification (e.g., Perez-Morales 1997; 
Nordenstam 1997a, 2003, 2007; Veldkamp & Lut 2009), and morphological 
treatments (e.g., Otieno & Tadesse 1992). 



Many of us have benefited from the Compositae Newsletter and it seems safe 
to say that without the efforts of Bertil Nordenstam and the support of the 
Swedish Museum the journal would not exist today. So, from the synantherology 
community, we say thank you to both. 



Comp. Newsl. 50,2012 




Fig. 1. Bertil Nordenstam collecting Compositae in Lesotho, close to the 
Oxbow lodge, above the Tsehlanyane River at ca. 2700 m. Photo: Vicki A. Funk. 



TIC A members are currently discussing the future of the Compositae Newsletter. 
So far everyone agrees that it should continue and most think that it will probably 
have to move to a mainly online journal perhaps similar to Phytokeys. No matter 
what is decided some means of support for the technical editing will have to be 
found. Also in this issue is a report on the TICA meeting in Montreal and it has 
additional information. Suggestions and members wishing to volunteer can email 
TICA ( synantherologist@gmail.com ). 

References 



Anderberg, A. A., Ghahremaninejad, F. & M. Kallersjo 2007. The enigmatic 
genus Dipterocome - Compositae. Comp. Newsl. 45: 23-36. 

Beltran, H. 1999. New combinations in Dendrophorbhim and Pentacalia 
(Senecioneae - Asteraceae) from Peru. Comp. Newsl. 34: 50-52. 

Beltran, H. & S. Baldeon 2009. A new species of Gynoxys (Asteraceae: 
Senecioneae) from Peru. Comp. Newsl. 47: 13-18. 



Comp.Newsl. 50,2012 



Bremer, K. 1988. A new corolla type from the Asteraceae-Arctotideae. 
Comp.Newsl. 15: 12-16. 

Freire, S. E., Sancho, G., Urtubey, E., Bayon, N., Katinas, L., Giuliano, D., 
Gutierrez, D., Saenz, A. A., Iharlegui, L., Monti, C. & G. Delucchi 

2005. Catalogue of Asteraceae of Chacoan Plain, Argentina. Comp. Newsl. 
43: 1-126. 

Herrera, P. P. & I. Ventosa 2005. Ecology of Cuban Asteraceae. Comp. Newsl. 
42: 89-108. 

Hind, D. J. N. & C. Jeffrey 2001. A checklist of the Compositae of Vol. IV 
of Humboldt, Bonpland & Kunth's Nova Genera et Species Plantarum. 
Comp. Newsl. 37: 1-84. 

Jansen, R. K., Palmer, J. D. & H. J. Michaels 1 988. Investigations of chloroplast 
DNA variation in the Asteraceae. Comp. Newsl. 15: 2-1 1 . 

Nordenstam, B. 1961. New species of Enryops . Bot. Notiser 114: 65-87. 

Nordenstam, B. 1988. Editorial. Comp. Newsl. 14: 1-2. 

Nordenstam, B. 1996. New combinations in Ecuadorian Senecioneae. Comp. 
Newsl. 29: 47-50. 

Nordenstam, B. 1997a. Nomenclatural notes on Ecuadorian Senecioneae. Comp. 
Newsl. 30: 46^19. 

Nordenstam, B. 1 997b. A new combination of Dendrophorbium (Compositae- 

Senecioneae). Comp. Newsl. 31: 22-23. 

Nordenstam, B. 1997. The genus Aequatorium B. Nord. (Compositae- 
Senecioneae) in Ecuador. Comp. Newsl. 31: 1-16. 

Nordenstam, B. 1999. New combinations in Monticalia (Compositae- 
Senecioneae) from Colombia. Comp. Newsl. 34: 29-36. 

Nordenstam, B. 2003. Alciope versus Capelio - a nomenclatural ordeal. Comp. 
Newsl. 39:48-51. 

Nordenstam, B. 2007. Senecio varicosus, a Linnaean name for the Balearic taxon 
known as Senecio rodriguezii (Compositae-Senecioneae). Comp. Newsl. 
45:8-15. 

Nordenstam, B. & P. B. Pelser 2005. Dauresia and Mesogramma: one new and 
one resurrected genus of the Asteraceae-Senecioneae from Southern Africa. 
Comp. Newsl. 42: 74-88. 

Nordenstam, B., Moussavi, M. & B. Djavadi 2002. A new annual species of 
Senecio from Iran. Comp. Newsl. 38: 42^16. 



Comp. Newsl. 50, 2012 



Nordenstam, B., Pelser, P. B. & L. E. Watson 2009. Lomanthus, a new genus 
of the Compositae-Senecioneae from Ecuador, Peru, Bolivia and Argentina. 
Comp. Newsl 47:33-40. 

Noroozi, J., Ajani, Y. & B. Nordenstam 2010. A new annual species of 
Senecio (Compositae-Senecioneae) from subnival zone of southern Iran 
with comments on phytogeographical aspects of the area. Comp. Newsl. 
48:43-62. 

Otieno, D. F. & M. Tadesse 1992. Pollen morphological studies in Senecio 
(Compositae-Senecioneae) from Ethiopia. Comp. Newsl. 20/21: 22-28. 

Pelser, P. B., Veldkamp, J.-F. & R. van der Meijden 2006. New combinations 
in Jacobaea Mill. (Asteraceae-Senecioneae). Comp. Newsl. 44: 1—11. 

Perez-Morales, C, Llamas Garca, F., Acedo Casado, C. & A. Penas 
Merino 1997. Typification and definition of Doronicum austriacum Jacq. 
(Asteraceae). Comp. Newsl. 30: 1-3. 

Veldkamp, J.-F. & C. Lut 2009. Senecio valerianifolius Wolf ex Link 
(Compositae), an enigma revealed. Comp. Newsl. 47: 4-7. 




Fig 2. Bertil Nordenstam collecting Lamprocephalus B. Nord. on the 
Waboomsberg, Western Cape. Photo: Gunilla Nordenstam 2006. 



Comp.Newsl. 50,2012 



News from The International Compositae 
Alliance 2012 Meeting 

Submitted by Vicki A. Funk & Luc Brouillet 

The 2012 TIC A (The International Compositae Alliance) meeting was held at the 
Montreal Botanical Garden in Montreal, Canada where it was hosted by Professor 
Luc Brouillet (MT). Although the number attending was lower than expected it 
was a young (mostly) and lively crowd with lots of discussion after most of the 
talks. Many interesting topics were presented, as you will see from the program 
and the abstracts that follow this brief message. 

During the business meeting we had an open discussion on TICA and there are a 
few important things to report. 

First, we took the opportunity to thank several of our members: 

Luc Brouillet, was thanked for organizing the meeting and field trip. 

Torsten Eriksson ( SBT) was recognized for his service as our Webmaster since 
TICA was started. Torsten and his family are moving to Bergen, Norway, for 
new jobs and he will no longer be able to manage the site. We deeply appreciate 
his efforts in helping TICA get started and his patience with questions from the 
members. Mauricio Diazgranados (MO) graciously agreed to take over as 
Webmaster and he will be sending out information to the membership in September 
when he starts his postdoctoral Fellowship at the Smithsonian Institution. 

Bertil Nordenstam (S) has been the editor of the Compositae Newsletter (CN) 
for many years and he has announced that he is stepping down after volume 50 
(this volume). Prof. Nordenstam was instrumental in changing CN into a proper 
journal and he pulled together each volume with the help of a Technical Editor. 
Many of us have benefited from the journal and we owe Prof. Nordenstam and 
The Swedish Museum of Natural History a debt of gratitude. We also thank the 
long-time Technical Editor Gunnel Wirenius Nohlin (starting with #24 in 
1994). 

Second, we decided to hold the next TIC Ameeting in conjunction with the XI Latin- 
American Botanical Association and the 64 th Congresso Nacional de Botanica do 
Brasil, to be held jointly in Salvador, Brazil in 2014. This will be the first time 
we have met jointly with such a large group and we are all looking forward to it. 
Our colleagues in Brazil and other Latin American countries are many and several 
have already volunteered to help. We plan on having one symposium during the 
Congress that will feature many of our Latin American Colleagues and the work 



10 Comp. Newsl. 50, 2012 



they have done on understanding the diversity in the family. After the congress 
we discussed having a one-day meeting of short papers to see what everyone is 
working on and to encourage collaborations. Nadia Roque has offered to organize 
a field trip to Chapada Diamantia after the meeting. 

Third, after a long discussion on the future of the Compositae Newsletter we 
decided that we should explore the possibility of publishing it electronically. Until 
now the journal has been supported by the Department of Phanerogamic Botany, 
Swedish Museum of Natural History, Stockholm, Sweden (S). Since 1988 they 
have provided a technical editor, and printed and mailed the journal to over 600 
scientists and libraries around the world, free of charge. Our colleagues at S have 
been more than generous for many years and we thank them very much. The 
members present felt that it was unlikely that we would find another institution 
to take over this responsibility. A suggestion to model the electronic version after 
Phytokeys was discussed and some follow up with the staff at Phytokeys has been 
positive but this would require page charges and/or dues. Right now the journal 
needs a new editor and nominations are encouraged. All opinions and offers of 
help are welcome; they will be compiled and circulated to T1CA members for 
comment ( synantherologist(5)gmail.com ). 

Finally, it with sadness that we mention that Abundio Sagastegui, one of our 
members, died recently. Below, his colleague Mike Dillon (F) has provided a few 
comments. 

Abundio Sagastegui Alva (11 July 1932 - 26 May 2012) 

Abundio Sagastegui Alva passed away on 26 May 2012 from complications due 
to an auto accident in Trujillo, Peru. Born in Guzmango, Cajamarca, Sagastegui 
rose to prominence within the Universidad Nacional de Trujillo (HUT) and 
later the Universidad Privada Antenor Orrego (HAO). He was one of the most 
influential and prolific botanists in Peru, having collected well over 18,000 
numbers and described nearly 90 species, mostly Asteraceae, including four new 
genera. No fewer than 30 species commemorate him, having been based upon his 
collections. His death is a great loss to Synantherology as well as to the botanical 
community as a whole. 



Comp.Newsl. 50,2012 



11 




MONTREAL, JULY 15-19, 2012 
PROGRAM 

All activities occur at the Biodiversity Centre, Jardin botanique de Montreal 

Sunday, 15 July 

17:00 to 19:00 - Biodiversity Centre Auditorium hall, Welcoming reception and 
registration 



Monday, 16 July 

08:30 registration 

09:00 Welcoming to TICA 2012 

09:15 Brouillet, L. - Hommage to Les Gottlieb 

09:30 Brouillet, L. - The Evolution of the Compositae: introduction to the 
symposium 

09:45 Scaglione, D. - Progress in genome sequencing of lettuce and other 
Cichorieae and Cardueae species 

10:30 Break 

11:00 PvEnaut, S. - The extent of genomic divergence among sunflower species 
with respect to their degree of geographic separation 

12:00-13:30 Lunch 

13:30 Noyes, R. - Apomixis in the Asteraceae: Still crazy after all these years 



12 Comp. Newsl. 50, 2012 



14:15 Diazgranados, M. - Frailejones (Espeletiinae Cuatrec): a recent rapid 
radiation shaped by the tropical Andes 

15:00 Discussion on the Evolution of the Compositae: the future 

1 7:00-2 1 :00 River Cruise and Banquet 

Tuesday, 17 July 

09:00 Smissen, R. - Using transcriptome data to test reticulate relationships among 
major clades of Gnaphalieae 

09:30 Cantley, J. T. Geoclimatic events of South America influence a west then 
northward progression of Neotropical Lepidaploinae (Vernonieae: Compositae) 

1 0:00 Schmidt-Lebuhn, A. - Testing the monophyly of Ozothamnus and Cassinia 

(Asteroideae: Gnaphalieae) 

10:30 Break 

1 1 :00 Funk, V. A.-Biogeographic patterns in Bidens native to Pacific Oceania 

11:30 Magee, A. - Systematics of southern African Anthemideae (Asteraceae): 
unraveling relationships within Pentziinae 

12:00-13:30 Lunch 

13:30 Bengtson, A. -- Phylogeny and evolution of the Metalasia clade 
(Gnaphalieae- Asteraceae) 

14:00 Garcia, F. - From Andean rainforests to high altitude Paramos: phylogenetic 
position, molecular dating and insights into the evolution and novel habits of 
lianescent and arborescent species of Pentacalia, Monticalia, Dendrophorbium 
and other relatives (Senecioneae) 

14:30 Poster session 

15:00 TICA meeting (informal discussion) -V. A. Funk 

Wednesday, 18 July 

09:00 Urbatsch, L. - Phylogeny of Ericameria and related genera of Astereae 
(Asteraceae) inferred from nuclear ribosomal and chloroplast sequence data 

09:30 Funk, V. A. - Systematics and Biogeography of the Liabeae (Compositae): 
an update 



Comp.Newsl. 50,2012 13 



10:00 Sancho, G. - A Phylogeny of the Gochnatieae: Understanding a critically 
placed tribe in the Compositae 

10:30 Break 

11:00 Keeley, S. - New Understanding of the Phylogeny and Biogeography of 
the Vernonieae 

11:30 Schmidt-Lebuhn, A. - Spatial diversity and collecting activity in the 
Australian native daisies (Asteraceae) 

12:00-12:30 Discussion 

12:30-13:30 Lunch 

13:30 Guided tour of the Montreal Botanical Garden 



Botanical Excursion 

Thursday, 19 July 

8:00 Departure for Vimy Ridge ultramafic formation 
Evening: Supper and lodging in Quebec City 

Friday, 20 July 

9:00 Departure for Chute-Panet bog and return to Montreal 



14 Comp. Newsl. 50, 2012 

The International Compositae Alliance (TICA) 
Montreal, Canada 15-19 July 2012 

ABSTRACTS 

Compiled and edited by: Luc Brouillet & Vicki A. Funk 

A Tribute to Les Gottlieb 

Luc Brouillet 



Centre sur la biodiversite, IRBV, Universite de Montreal, 4101 Sherbrooke St E, 

Montreal, QC Canada H1X 2B2 

luc.brouillet@umontreal.ca 

The TICA 2012 meeting is dedicated to the memory of Dr. Les Gottlieb, a life- 
long student of evolution using Stephanomeria (Compositae, Cichorieae) and 
Gilia (Polemoniaceae) as models. Born in New York City, he finished his Ph. D. 
at the University of Michigan in 1969; his topic was speciation in Stephanomeria. 
He spent his entire career as a plant evolutionary biologist at the University of 
California-Davis. His early studies focused on the process of speciation using 
Stephanomeria as a model system, where he documented homoploid hybrid 
speciation, polyploid speciation, and sympatric speciation, using isozyme 
data. The last case was particularly interesting since it involves the diploid 
S. malheiirensis Gottlieb, which originated within a population of its progenitor, 
S. exigua subsp. coronaria. Dr. Gottlieb made an outstanding demonstration of 
this speciation event, investigating numerous aspects of the phenomenon. 
In addition he published many papers on fundamental issues in evolution and 
systematics. More recently, he contributed to phylogenetic studies within the tribe 
Cichorieae in western North America, and he wrote Stephanomeria and relatives 
for the Flora of North America. Dr. Gottlieb was an outstanding example of 
an evolutionary scientist who used Compositae as a model system, while 
contributing to the systematics of the group and so inspired a generation of young 
systematists to think more broadly and to always strive to understand the context 
of the morphology they were documenting. 



Comp. Newsl. 50, 2012 15 

Symposium: Compositae Evolution 

(Alphabetical by Author) 
The Evolution of Compositae: an introduction 

Luc Brouillet 



Centre sur la biodiversite, IRBV, Universite de Montreal, 4101 Sherbrooke St E, 

Montreal, QC Canada H1X 2B2 

luc.brouillet@umontreal.ca 

The study of the evolution of Compositae has two facets: the study of the evolution 
of Compositae groups and the study of evolution using Compositae as model 
organisms. In recent years, while Comps systematists were busy developing a 
global phylogeny and biogeography of the family, other researchers were making 
progress in several fields related to their evolution: paleobotany, development, 
phylogeography, reproduction, speciation and genomics. The need of dated fossils 
to calibrate Compositae phylogenies has been partly filled recently by the discovery 
of fossil mutisioid-carduoid heads from the Eocene of Patagonia (Barreda et al. 
2010), as well as pollen of the same age from the southern hemisphere, dating the 
origin of Comps to about 50 ma. New insights into the origins of the capitulum 
were provided by a developmental study of Calyceraceae inflorescences, showing 
the cymose origin of the Comp head (Pozner et al. 2012). On the developmental 
genetics front, a study of the genes controlling head and floret development in 
Comps since the late 1990s has resulted in the development of a model of head 
development in Comps (Laitinen et al. 2006), as well as in comparative studies. 
The Compositae are also known for their sporophytic self-incompatibility system, 
the genetics of which has started receiving attention (e.g. Hiscock et al. 2003), 
a major development given its consequences for the evolution of species in the 
group. Another reproductive system studied well is agamospermy (apomixis), 
which has been examined in many taxa from genetic and evolutionary standpoints. 
Phylogeographic studies have been relatively scarce in the family but have 
provided insights into the evolution of some species or species groups, notably 
in Senecio and relatives. The study of speciation and its genetic aftermaths has 
been the object of in-depth studies in many genera, most notably in Helianthus 
(Rieseberg lab) and in Tragopogon (Soltis lab), including molecular, cytological 
and genomic approaches. In the genomics field, a new genome size database was 
developed (http://www.asteraceaegenomesize.com/) for the family. But the main 



16 Comp. Newsl. 50, 2012 



recent progress has been the complete sequencing of the chloroplast genome of 
lettuce and sunflower (Timme et al. 2007), as well as the recent sequencing of 
the nuclear genome of these two organisms (Michelmore and Rjeseberg labs), 
which will soon be made available to the community. Genomics has already 
shown that the Compositae genome was subjected to multiple polyploidization 
events (Barker et al. 2008). These new developments in Compositae research will 
open the door to new venues of research and new tools for the students of Comp 
phylogeny, and reciprocally, evolutionary biologists will benefit from better and 
more detailed phylogenies. 

Barreda, V. D. et al. 2010. Eocene Patagonia fossils of the Daisy family. Science 
329: 1621. 

Pozner, R. et al. 2012. Evolutionary origin of the Asteraceae capitulum: Insights 
from Calyceraceae. Amen J. Bot. 99: 10-13. 

Laitinen, R. A. E. et al. 2006. Patterns of MADS-box gene expression mark 
flower-type development in Gerbera hybrida (Asteraceae). BMC Plant Biology 
6: ll.doi:10.1186/1471-2229-6-ll 

Hiscock, S.J. et al. 2003. Sporophytic self-incompatibility in Senecio squalidiis 
L. (Asteraceae) - the search for S. J. Expermental Bot. 54: 169-174. doi: 10.1093/ 
jxb/erg005 

Timme, R. E. et al. 2007. A comparative analysis of the Lactuca and Helianthus 
(Asteraceae) plastid genomes: Identification of divergent regions and categorization 
of shared repeats. Amen J. Bot. 94: 302-312. 

Barker, M. S. et al. 2008. Multiple paleopolyploidizations during the evolution of 
the Compositae reveal parallel patterns of duplicate gene retention after millions 
of years. Mol. Bio. Evol. 25: 2445-2455. doi: 10.1093/molbev/msnl87 



Comp. Newsl. 50, 2012 17 

Frailejones (Espeletiinae Cuatrec): a recent rapid radiation 
shaped by the tropical Andes 

Mauricio Diazgranados 1,2 & Janet C. Barber 1 



'Department of Biology, Saint Louis University, 3507 Laclede Ave., 

St. Louis, MO 63 103 USA 

2 Missouri Botanical Garden, 4344 Shaw Boulevard, 

Saint Louis, MO 63110 USA 

espeletias@gmail.com 

The paramo ecosystem, located above the tree line in the tropical Andes, has been 
the setting for some of the most dramatic recent rapid plant radiations. With an 
estimated age of 2-4 million years, the paramo is the world's most diverse high- 
elevation ecosystem. Today 141+ species of frailejones (subtribe Espeletiinae 
Cuatrec, Asteraceae) dominate this ecosystem. Frailejones have intrigued 
naturalists and botanists, not just for their appealing beauty and impressive 
morphological diversity, but also for their remarkable adaptations to the extremely 
harsh environmental conditions of the paramo. The most recent classification of 
the subtribe has eight genera (Carramboa, Coespeletia, Espeletia, Espeletiopsis, 
Libanothamnus, Paramiflos, Ruilopezia and Tamania). Previous attempts to 
reconstruct the evolutionary history of this group failed to resolve relationships 
among genera and species, and there is no agreement regarding the classification 
of the group. For this study, sequence data included nrDNA (ITS and ETS) and 
cpDNA (rpl 16), for a total of 2954 aligned bp. Fragment analysis was performed 
with AFLP data using 28 primer combinations and yielding 1665 fragments. 
Phylogenies reconstructed suggest that various clades are monophyletic, but 
the phylogenetic signal may be misled by hybridization and incomplete lineage 
sorting. The origin of the subtribe in Venezuela is supported. A shallow phylogeny 
with short branches and the impressive morphological diversity suggest a recent 
rapid radiation. A Monte Carlo permutation test shows a very strong geographic 
structure in the phylogeny. Venezuelan paramos show more phylogenetic 
overdispersion, whereas Colombian paramos generally exhibit phylogenetic 
clustering, with sympatric species closely related to each other. 



Comp. Newsl. 50, 2012 



Apomixis in the Asteraceae: Still crazy after all these years 



Rick Noyes 



University of Central Arkansas, Dept. of Biology, 180 Lewis Science Center, 

Conway, Arkansas 72035, USA 

moyes@uca.edu 

The Asteraceae is commonly listed as one of the principal families within which 
asexual reproduction by seed, i.e., apomixis, is prolific. Critical review of the 
literature indicates that naturally occurring apomixis is robustly indicated for 22 
genera in seven tribes of Asteraceae, all but one of which occurs in subfamily 
Asteroideae. Consideration of 45 additional reports indicates that apomixis for 
30 genera is contra-indicated, documenting possible developmental abnormalities 
in otherwise sexual taxa. Data are strongly indicative or equivocal for effective 
apomixis for an additional fifteen genera, but thorough documentation is wanting. 
Thus our state of knowledge of apomixis in the Asteraceae is generally poor. 
Apomixis has been extensively investigated for three genera of Asteraceae: 
Erigeron, Hieracium (Pilosella), and Taraxacum. Evidence from crossing and 
developmental studies in these taxa indicates that apomixis in the Asteraceae 
is generally facultative, resulting from the occasional production of reduced 
megagametophytes that can participate in biparental reproduction. For instance, 
for Erigeron annum, data indicates that individual plants, on average, produce >2% 
ovules that undergo meiotic rather than apomeiotic division of the megasporocyte. 
Strict apomictic development occurs, but is rare. From a biosystematic perspective, 
some of the biggest challenges in apomixis biology include 1) understanding why 
apomixis occurs in some taxa but not in others, 2) interpreting the number of 
unique origins of apomixis within complexes, 3) elucidating the environmental 
and demographic conditions that favor the evolution of apomixis in populations, 
and, 4) determining how the genes for apomixis can spread to sexual populations 
by pollen. 



Comp.Newsl. 50,2012 19 

The extent of genomic divergence among sunflower species 
with respect to their degree of geographic separation 

SEBASTIEN RENAUT & LOREN H. RlESEBERG 



Department of Botany, University of British Columbia, 6270 University Blvd, 

Vancouver, BC, V6T 1Z4, Canada 

sebastien. ernaut@gmai 1 . com 

Levels of differentiation among populations can be highly variable across the 
genome. During allopatric speciation, divergence should accumulate across 
the genome due to the action of both drift and selection. In contrast, during 
speciation with gene flow, accentuated divergence should be restricted to loci 
under divergent natural selection. Here, we report on high resolution genomic 
scans of differentiation among several pairs of sunflowers {Helianthus spp.) taxa 
that vary in their divergence time and degree of geographic separation. We find 
that in all comparisons, genomic regions of divergence are numerous and small 
(< 1 centiMorgan). In addition, the proportion of divergent loci fixed by selection 
was higher in sympatric (43%) than allopatric species pairs (24%), thus confirming 
the more prominent effect of selection in shaping genomic divergence in sympatry. 
Lastly, we also find that among independent species pairs, patterns of genomic 
divergence are surprisingly repeatable, especially in highly differentiated regions. 
This is due, at least in part, to a repeatable heterogeneous pattern of recombination 
rates along the genome in independent species pairs. In conclusion, the genomic 
clustering of highly divergent loci is not influenced by the extent of ongoing gene 
flow, but probably by other factors such as recombination rates. 



20 Comp.Newsl. 50,2012 

Progress in genome sequencing of lettuce and other Cichorieae 
and Cardueae species 



Davide Scaglione 1 , Sebastian Reyes-Chin-Wo 1 , Zhiwen Wang 2 , 

Christopher Beitel 1 , Alexander Kozik 1 , Song Chi 2 , Wenbin Chen 2 , 

Maria Jose Truco 1 , Xun Xu 2 , Lutz Froemcke 1 , Dean Lavelle 1 , 

Bicheng Yang 2 , Ian Korf 1 , Jun Wang 2 & Richard Michelmore 1 



'Genome Center, University of California, Davis, CA, USA 

2 BGI, Shenzhen, China 

gianza@hotmail.it 

The Compositae Genome Project (http://compgenomics.ucdavis.edu) is a 
collaborative project to analyze genetic diversity in the Compositae plant family. 
Lettuce, an economically important member of this family, is being studied to 
correlate genotype with phenotypic variation in domestication and agriculturally 
significant traits. Several resources have been generated including lettuce 
transcriptome, gene-space, and whole genome assemblies. The whole genome 
of Lactuca sativa cv. Salinas has been sequenced in collaboration with the BGI 
and a consortium of ten breeding companies. From ~70x coverage with high 
quality, filtered Illumina reads, 2.5 Gb (93% of the entire genome) was assembled 
into scaffolds with a N50 of 461,580 bp. Over 11,000 loci from an ultra-dense, 
transcript-based genetic map was used to assess the quality of the assembly. Over 
95% of 3,100 scaffolds that had multiple unigenes were genetically validated 
and could be ordered in chromosomal linkage groups. The resulting data are 
displayed using GBrowse. More scaffolds are being assigned to chromosomal 
linkage groups using genotyping by sequencing of 99 RILs from the reference 
L. sativa cv. Salinas x L. serriola mapping population. Chromosomal orders 
are being refined using population and syntenic information. Approximately, 
45,000 gene models have been predicted using several automated annotation 
pipelines. Manual curation of these models is underway to refine the annotations. 
The genome sequence is being used to clone and functionally validate genes for 
disease resistance and development. Generation of ultra-dense maps of chicory 
and artichoke has also been initiated. 



Comp. Newsl. 50, 2012 21 

General Papers 

(Alphabetical by Author) 

Phylogeny and evolution of the Metalasia clade 
(Gnaphalieae-Asteraceae) 

Annika Bengtson l , Arne Anderberg 2 & Per Ola Karis 1 



'Department of Botany, Stockholm University, SE-106 91 Stockholm 

2 Department of Phanerogamic Botany 

Swedish Museum of Natural History, Box 50007, SE-104 05 Stockholm 

annika. bengtson@botan.su 

Metalasia is an endemic South African genus of ericoid shrublets in the tribe 
Gnaphalieae, currently consisting of 57 species with their main distribution in 
the Cape Floristic Region. Like many other CFR plant clades most species of 
Metalasia have very narrow distributions whereas a few are found over larger 
areas. Earlier studies based on morphological data alone have tried to disentangle 
the relationships between Metalasia and its allies, but until the present project 
was initiated, no analyses of molecular data had been performed. Our initial study 
based on DNA sequence data gave support for a monophyletic Metalasia clade 
comprising eight genera and indicating that some generic realignments may be 
necessary. The focus of the subsequent work has been a larger study including all 
currently recognized species of Metalasia, utilizing a combination of molecular 
and morphological data. In contrast to the prevailing cladistic hypothesis from 
morphological data, our phylogeny show that the species of Metalasia form two 
equally sized, well-supported evolutionary lineages with different distribution, 
and that Metalasia in its current sense is paraphyletic. Biogeographical analyses 
will reveal more about the evolutionary history of the Metalasia clade. 



22 Comp.Newsl. 50,2012 

Geoclimatic events of South America influence a west 

then northward progression of Neotropical Lepidaploinae 

(Vernonieae: Compositae) 

Jason Tyler Cantley & Sterling Keeley 



University of Hawaii at Manoa, 3190 Maile Way, St John 101, 

Honolulu, Hawaii 96822, USA 

jaecan@gmail.com 

The historical biogeography of the open canopy dry habitats that cover much 
of South America remains largely unexplored despite detailed reconstructions 
of paleogeology and paleoclimatology. To test the floristic biogeographical 
relationships of these habitats the systematic and biogeographical relationships 
in the Lepidaploinae were investigated. The Neotropical Lepidaploinae is the 
largest and most broadly distributed subtribe of tribe Vernonieae: Compositae. 
A molecular phylogeny was constructed for 91 species of the Lepidaploinae 
using three chloroplast DNA regions (frnLc-f, ndhF, matK) and the nuclear 
ITS region. Biogeographic and molecular clock analyses indicate that the 
Lepidaploinae originated on the Brazilian Shield about 20-12.5 mya. The 
subsequent diversification of five major lineages is closely tied to geologic and 
climatic events in South America such as the Andean uplift, periodic lowland 
marine/freshwater incursions and the development of the Amazon Rainforest as 
an exclusionary habitat barrier. As new suitable habitat was formed on a changing 
South American continent, Lepidaploinae lineages advanced westward from the 
Brazilian Shield to the Andes; and then northward, following the uplift of the 
Andes Mountains. The timing of diversification of Lessingicmthus s.s. in Cerrado 
habitat is correlated to the worldwide rise in C4 grasses. Lepidaploa s.s. lineage 
reached Central America and the West Indies within the last four million years, 
coincident with the closure of the Isthmus of Panama and long distance dispersals, 
respectively. Overall, the lineage diversifications of subtribe Lepidaploinae may 
indicate how South America's complex paleogeoclimatic history have influenced 
the evolution of Lepidaploinae as well as other angiosperm lineages. 



Comp.Newsl. 50,2012 23 

Biogeographic patterns in Bidens native to Pacific Oceania 



Vicki A. Funk 1 , Gabriel Johnson 2 , Matt Knope 3 , Mauricio Bonifacino 4 , 
Daniel Crawford 5 , Fred Ganders 6 , Dave Lorence 7 , Jean-Yves Meyer 8 & 

Warren L. Wagner 1 



'Smithsonian Institution, National Museum of Natural History, 

Department of Botany, Washington, DC, USA 

2 Smithsonian Institution, National Museum of Natural History, 

Laboratory of Analytical Biology, Washington, DC, USA 

Stanford University, Palo Alto, CA, USA 

"Universidad de la Republica, Montevideo, Uruguay 

-University of Kansas, Lawrence, USA 

6 University of British Columbia, Vancouver, Canada 

7 National Tropical Botanical Garden, Kalaheo, HI, USA 

Government of French Polynesia, Research Department, Papeete, Tahiti 

funkv@si.edu 

The genus Bidens (Compositae or Asteraceae: Coreopsideae) has over 200 species 
and includes 37 species in Pacific Oceania. Most species are from Hawaii or 
French Polynesia but a few other species are scattered in other parts of the Pacific 
region. Over one hundred samples from the Bidens and related genera of the 
Coreopsideae have been chosen for testing using specific nuclear and chloroplast 
markers. Specifically, ITS, ETS, r/?/132trnL, psbA, and trnQrps6 regions were 
used. Do all of the Bidens found in the Pacific region form a monophyletic group? 
Yes and No. All species from the Hawaiian Islands, the Marquesas, and the Society 
Islands form a monophyletic group although relationships among the species 
found on each islands are still under investigation. But species from Australia, 
Starbuck Island, and Socorro Island are from independent lineages within the 
genus. Where are the closest relatives of the Bidens in the Pacific region? Previous 
work indicated that Bidens from western North America might be the closest 
relatives for the Hawaiian species and this is supported by this research. How 
does the biogeography of Bidens compare to other members of the Compositae 
in Pacific Oceania? Of the 164 species of Compositae native to Pacific Oceania, 
Bidens has the most diverse distribution and is one of the four lineages whose 
ancestry can be traced to Western North America. 



24 Comp. Newsl. 50, 2012 

Systematics and Biogeography of the Liabeae (Compositae): 

an update 

Vicki A. Funk, Carol Kelloff & Raymund Chan 



Smithsonian Institution, National Museum of Natural History, 

Department of Botany, Washington, DC, USA 

funkv@si.edu 

The tribe Liabeae (Compositae) contains ca. 175 species distributed in 18 genera 
and its members occupy a variety of habitats in Andes Mountains of South America 
as well as Mexico, Central America, and the West Indies. The tribe is characterized 
by a combination of morphological characters, including opposite leaves with 
white-tomentose pubescence beneath and often with venation strongly trinervate, 
yellow ray and disk florets, oblong or columnar achenes usually with a biseriate 
pappus that frequently consists of outer scales and inner scabrous bristles, and the 
frequent occurrence of latex. DNA sequence data from the nuclear ribosomal ITS 
region and four chloroplast regions (trnL-F, 3' end of ndhF, matK, psbA; a total 
of more than five kb of sequence data) were used to infer a phylogeny. The data 
were analyzed using Maximum parsimony, Maximum likelihood, and Bayesian 
inference posterior probabilities. The results support the monophyly of the tribe 
and show a consistent placement for all genera except Cacosmia (3 species). Four 
well-supported clades are recovered in the remainder of the tribe, all recognized as 
subtribes. Liabineae are the sister group of the rest of the tribe. Sinclairineae are the 
sister group of Munnoziinae plus Paranepheliinae. The genus Bishopanthus could 
not be confidently placed in any of the subtribes; molecular study is not possible. 
The phylogeny slightly alters the previous assumptions about the biogeography 
and it seems that the Liabeae originated in the Central and Northern Andes and 
spread north and south with several independent introductions into Mexico and 
Central America and one into the Caribbean. With the exception of the Liabeae 
(Andes) and Moquineae (Brazil), all of the tribes in the subfamily Cichoroideae 
are either restricted to or have their basal grade in Africa. 



Comp. Newsl. 50, 2012 25 

From Andean rainforests to high altitude Paramos: 

phylogenetic position, molecular dating and insights into 

the evolution and novel habits of lianescent and arborescent 

species of Pentacalia, Monticalia, Dendrophorbium and other 

woody relatives (Senecioneae) 

Frederico Garcia 



Universidad INCCA de Colombia, Carrera 13 # 24-15, 

Bogota, Colombia 

fgarciay@gmail.com 

The high Andes of South America provide a number of geographically and 
ecologically isolated systems inhabited by plants showing highly modified 
morphological characters. In this study, sequences from the ITS region of nuclear 
ribosomal DNA are used to test previous hypotheses about the phylogenetic 
position of divergent Andean genera, classified in subtribes Tussilagininae and 
Senecionineae. Gene phylogenies based on maximum parsimony and maximum 
likelihood reveal relationships of arborescent Andean Tussilagininae like Gynoxys, 
Paragynoxys and Aequatorium to Central and North American Tussilagininae 
(Pittocaulon, Robins onecio), whereas a genus (Acrisione) appears related to a 
New Zealand clade. We also analyzed the controversial definition of the scandent 
species of the genus Pentacalia in relation to its segregates of erect, high altitude 
species of Monticalia sensu C. Jeffrey. The purpose was to test the hypothesis 
of a high altitude origin of scandent forms (i.e. Pentacalia species) from erect 
Monticalioid ancestors, with an ulterior process of etiolation, all this resulted 
in a huge radiation of vine species. Arborescent species of Dendrophorbium 
appear as ancestral forms of another independent vine clade. Viny taxa (e.g. 
Cabreriella, Pentacalia cuatrecasana, Pentacalia rugosa) from Sierra Nevada 
e.g. Santa Marta (a Cretaceous mountain range isolated from the Andes) appear in 
different clades, but always in an ancestral position related to core representatives 
of each clade. Also presented is a phylogenetic dating of Senecioneae based on 
parametric methods (Multidivtime) and six fossil references. The results suggest 
a vicariant event separating the three clades of the tribe by the end of upper 
Miocene, coinciding with Miocene Climatic Optimum. Divergence of South 
American Senecioneae is discussed in the context of Middle Miocene Caribbean 
and Andean tectonics. 



26 Comp. Newsl. 50, 2012 



New Understanding of the Phylogeny and Biogeography of 

the Vernonieae 



Sterling Keeley 



University of Hawaii at Manoa, 3190 Maile Way, St John 101, 

Honolulu, Hawaii 96822, USA 

sterling@hawaii.edu 

The tribe Vernonieae is among the largest in the Compositae with ca. 1500 
taxa. There are two centers of diversity, southern Brazil in the New World and 
southeastern Africa in the Old World. The previous molecular phylogeny indicated 
an African/Madagascan origin for the tribe with a basal grade of Old World taxa 
from which New World species arose. Subtribal/generic relationships within and 
between hemispheres remained unclear, however. A new phylogeny based on 
chloroplast and nuclear DNA sequence data of over 300 taxa provides insight into 
subtribal relationships and dispersal pathways. Africa is confirmed as the ancestral 
region for the extant members of the tribe and the original source of both New 
and Old World lineages. Australian and all but one lineage from Southeast Asia 
(not including India) are of the result of dispersals from African ancestors and 
suggest a possible route to Hawaii. There was at least one back dispersal from the 
New to the Old World giving rise to a clade of Southeast Asian taxa. New World 
Vernonieae are monophyletic and apparently the result of a single dispersal from 
Africa to Brazil. Within the Americas movement can be traced from southeastern 
Brazil, along the Andes, to Central America and Mexico and the West Indies in 
several clades including the largest subtribe, the Lepidaploinae. Lineage diversity 
is limited in eastern North America. Subtribal and generic delimitations will need 
revision especially in the Old World Erlangeinae, Centrapalinae and Linziinae and 
the New World Lepidaploinae and Piptocarphinae. 



Comp.Newsl. 50,2012 27 

Systematics of southern African Anthemideae (Asteraceae): 
unraveling relationships within Pentziinae 

Anthony Magee 1,2 , A. N. Nicolas 3 , P. M. Tilney 2 & G. M. Plunkett 3 

'Compton Herbarium, Biosystematics and collections, 

South African National Biodiversity Institute, Private Bag X7, 

Claremont 7735, Cape Town, South Africa 

department of Botany and Plant Biotechnology, University of Johannesburg, 

P.O. Box 524, Auckland Park 2006, Johannesburg, South Africa 

'Cullman Program for Molecular Systematics, The New York Botanical Garden, 

2900 Southern Blvd., Bronx, NY 10458-5126, USA 

A.Magee@sanbi.org.za 

The southern African taxa comprise the earliest diverging lineages within the 
largely northern hemisphere tribe Anthemideae. Improved understanding of the 
evolution, diversification and biogeographical history of these southern African 
lineages is therefore crucial. Generic and species delimitations are, however, far 
from satisfactory. As a result taxonomic and phylogenetic investigations need 
to be conducted concurrently. The interesting phylogeographical link between 
the Pentziinae and the northern hemisphere subtribes and the disjunct north- 
south distribution of Pentzia made this subtribe a clear choice as the starting 
point for broader systematic studies of southern African Anthemideae. The 
Pentziinae comprise seven almost-exclusively southern African endemic genera 
(Cymbopappus, Foveolina, Marasmodes, Myxopappus, Oncosiphon, Pentzia and 
Rennera) and ca. 59 species. Several of the genera in the subtribe were previously 
united under Pentzia. Despite this there remains doubt regarding the monophyly 
of Pentzia, particularly in relation to Cymbopappus and Marasmodes. It is clear 
that a re-assessment of generic delimitations within the subtribe should be coupled 
with a comprehensive taxonomic revision of the central genus Pentzia. The 
genus comprises ca. 27 species and has not been revised since the now outdated 
synopsis of Hutchinson in 1917. Generic delimitations and relationships within 
the Pentziinae are explored using morphological, anatomical and molecular 
sequence data (ndhF, psbA-trnH, r/?/32-trnL, nrlTS). Phylogenetic analyses for 
71 accessions (49 species) representing 87% of the subtribe indicates that several 
of the genera are not monophyletic as currently circumscribed. This is further 
corroborated by morphological and anatomical characters. 



28 Comp. Newsl. 50, 2012 

A phylogeny of the Gochnatieae: Understanding a critically 
placed tribe in the Compositae 



Vicki A. Funk 1 , Gisela Sancho 2 , Nadia Roque 3 , Carol L. Kelloff 1 , 
Iralys Ventosa-Rodrigues 4 , Raymund Chan 1 , & Mauricio Bonifacino 5 



'Smithsonian Institution, National Museum of Natural History, 

Department of Botany, Washington, DC, USA 

2 Museo de La Plata, Paseo del Bosque s/n, 1900 La Plata, BA, Argentina 

3 Instituto de Biologia, Universidade Federal da Bahia, 

Campus Universitario de Ondina, 40170-110 Salvador, Bahia, Brazil 

4 Pasaje El Astillero No. 8, (4401 ) Vasqueros, Salta, Argentina 

5 Laboratorio de Botanica, Facultad de Agronomia, Montevideo, Uruguay 

funkv@si.edu 

The Gochnatioideae clade is basal to most Asteraceae, excluding the 
Barnadesioideae, Stifftioideae, Mutisioideae, and Wunderlichioideae clades. 
Gochnatieae, the only tribe of Gochnatioideae comprises four genera: 
Cnicothamnus , Cyclolepis, Gochnatia, and Richterago. The recognition of 
Gochnatieae was the result of resolving the polyphyly of the Mutisieae suggested 
on the base of morphology. The tribe Gochnatieae can be defined by a combination 
of morphological characters, although these features are present in other basal 
clades. By far, Gochnatia, which traditionally includes about 70 species, is the most 
complex of the tribe and a key taxon to understand Gochnatieae. Within Gochnatia, 
some groups of species, treated as sections, are morphologically well defined and 
identifiable. Recently, some of the traditional sections of Gochnatia have been 
treated at the genus rank. However, until the present study, a comprehensive and 
complete phylogenetic analysis including most of its representative species has 
not being carried out. Our results suggest a paraphyletic Gochnatia that is here 
resolved by the circumscription of segregate genera, some of which are already 
established and other of which are new. Some morphological characters support 
these new genera and allow us to present a new comprehensive classification of 
Gochnatieae. 



Comp.Newsl. 50, 2012 29 

Testing the monophyly of Ozothamnus and Cassinia 
(Asteroideae: Gnaphalieae) 

Alexander Schmidt-Lebuhn & Lee Constable 



CSIRO Plant Industry / Centre for Australian National Biodiversity Research, 

Clunies Ross Street, 

Canberra, ACT 2601, Australia 

alexander.schmidt-lebuhn@csiro.au 

The Australian and New Zealand shrubby everlastings of the genera Ozothamnus 
and Cassinia and their small satellites represent about 100 species or ca. 10% 
of the Australian daisy flora. Generic delimitations in the group have long been 
controversial and subject to revision. In their current circumscription, the two 
large genera are differentiated based on the presence or absence of paleae and the 
shape of the phyllaries, although several exceptions exist. It is suspected that at 
least more heterogeneous Ozothamnus is non-monophyletic, and that several of 
the smaller genera in the group may constitute apomorphic segregates (defined 
merely by loss of the pappus, etc.), but no comprehensive phylogenetic study 
had been undertaken. We have produced a molecular phylogeny based on very 
broad sampling and using nuclear ITS, ETS and chloroplast psbA-trnH, matK- 
psbA and ycf6-psbM regions. We submitted sequences from the two main genera 
to Templeton and K-H-Rell tests to compare phylogenies from constrained 
analyses statistically against the best trees from unconstrained analysis. We found 
Ozothamnus to be non-monophyletic even in a very narrow circumscription, 
i.e. after the removal of section Hebelaena and several other divergent species. 
Cassinia, on the other hand, is likely a natural group. Nuclear data suggest that 
Calomeria, Cassinia, Hackeria and Odixia are nested in Ozothamnus; chloroplast 
data provide poorer resolution and a somewhat contradictory topology but the 
results of the constraint tests are identical for both datasets. 



30 Comp.Newsl. 50,2012 

Spatial diversity and collecting activity in the Australian native 

daisies (Asteraceae) 



Alexander Schmidt-Lebuhn, Nunzio J. Knerr 
& Carlos E. Gonzalez-Orozco 



CSIRO Plant Industry / Centre for Australian National Biodiversity Research, 

Clunies Ross Street, Canberra, ACT 2601, Australia 

alexander.schmidt-lebuhn@csiro.au 

Australia is home to ca. 1,000 currently recognized species of native Asteraceae, 
50% of them Gnaphalieae, 30% Astereae, 10% Senecioneae. We conducted spatial 
analyses at the 1° scale to examine the distribution of species richness, collecting 
activity and various measures of phylogenetic diversity. In a first step, we queried 
all Asteraceae specimen data from Australia's Virtual Herbarium, removed 
duplicates and deleted all specimens lacking point data and all geographic outliers. 
The final dataset comprised 114,537 collections of 968 species. Three different 
approaches of inferring species diversity were compared; the results were broadly 
equivalent, with the twenty hotspots found in the south-east of the continent and 
Tasmania in all cases. Reliance on documented species numbers is direct but leads 
to a distorted perception of the relative local diversity due to very uneven collecting 
efforts. The Chao 1 estimator of species richness corrects those distortions but 
is unavailable for about 50% of Australia due to insufficient local collections. 
Finally, a distribution modelling approach provides inferences for all of Australia 
but overestimates species numbers across the board. To obtain branch lengths for 
phylogenetic diversity, a molecular phylogeny (\TS,psbA-trnH, matK, trnL-trnF) 
was generated with 159 of the 164 genera as OTUs. Phylogenetic diversity was 
inferred to be high in the south-west and south-east, and phylogenetic endemism 
to be high in the tropical north, the south-west, Tasmania and in border ranges 
between Queensland and New South Wales. 



Comp. Newsl. 50, 2012 31 

Using transcriptome data to test reticulate relationships 
among major clades of Gnaphalieae 

Rob Smissen, Merce Galbany-Casals & Ilse Breitwieser 



Landcare Research, New Zealand, PO Box 40, Lincoln, 

Canterbury 7640, New Zealand 

smissenr@landcareresearch.co.nz 



Previously we have postulated ancient allopolyploid origins for several diverse 
and widely distributed clades of Compositae tribe Gnaphalieae on the basis of low 
copy number nuclear gene phylogenies. Presently we are making use of massively- 
parallel sequencing technology to further test and elaborate our hypotheses. Using 
the Roche GS-Junior platform we have generated between 38 and 52 Megabases 
of sequence for four species of Gnaphalieae, two putative allotetraploids and two 
putative diploids. We are employing two approaches to incorporate these data into 
our research. Firstly, we are using assembled cDNA sequences to characterise 
additional genes for amplification by PCR and phylogenetic analysis. Secondly, we 
are attempting to identify orthologous sequences from the transcriptome assemblies 
and conduct phylogenetic analyses of these directly. Although preliminary results 
are consistent with our hypothesised allopolyploid relationships, at the level of 
phylogenetic divergence across the tribe, both approaches are proving difficult 
to implement. As is the case in most phylogenetic studies using nuclear genes, 
establishing orthology/paralogy relationships among sequences is a major 
problem. This talk will present an update of evidence suggesting the importance 
of allopolyploidy in the phylogeny of the tribe Gnaphalieae and discuss challenges 
posed by phylogenetic analysis of genome scale data sets. 



32 Comp. Newsl. 50, 2012 



The ycfl and other chloroplast gene regions for phylogenetic 
assessment of tribe Astereae 



Lowell Urbatsch 1 , Lauren Eserman 1 , Kurt Neubig 2 , Roland Roberts 3 

& Ross Holston 1 



'Louisiana State University, Baton Rouge, LA, USA 

2 University of Florida, Gainesville, FL, USA 

3 Towson State University, & Towson, MD, USA 

leu@lsu.edu 

Species centered about Ericamerid and their derivatives are abundant in 
western North America especially in the California Floristic Province, the Great 
Basin, and Rocky Mountains. The ycfl chloroplast region was sequenced in 
approximately 100 representative taxa. Datasets were aligned using T-Coffee and 
manually adjusted. Variable and parsimony informative sites were obtained using 
MEGA 5. Models were chosen for each using the AIC criterion implemented in 
jModelTest, and all analyses were run using the GTR model. The analyses were 
run using MrBayes on the CIPRES portal. Trees represent a 50% majority-rule 
consensus tree of post-burnin trees from 4 independent runs constructed using 
the sumt command in MrBayes. The great success story in these investigations 
has been Ericameria. Various workers have shown it to be comprised of species 
traditionally accommodated in Chrysothamnus, Macronema, Haplopappus sect. 
Asiris, and Stenotopsis. ITS, ETS, and chloroplast markers robustly support this 
clade. Chrysothamnus, a long recognized genus, is not supported by any of the data 
sets. Comparisons between the ITS/ETS and chloroplast regions among the other 
entities investigated will be made. Data from ycfl sequences of basal Astereae and 
taxa from different geographic regions suggests that it is an appropriate candidate 
DNA region for further testing the phylogeny for tribe Astereae. 



Comp. Newsl. 50, 2012 33 

Posters 

(Alphabetical by Author) 

Taxonomic revision of Baccharis subgen. Tarchonanthoides 

(Compositae: Astereae): a group from the South American 

grasslands and savannahs 

Gustavo Heiden & Jose Rubens Pirani 



Laboratorio de Sistematica Vegetal, Departamento de Botanica, 

Instituto de Biociencias, Universidade de Sao Paulo, 

Rua do Matao Travessa 14, 321, 05508-090, Sao Paulo, SP, Brazil 

gustavo.heiden@gmail.com 

Baccharis L. is a New World genus with about 400 species and broadly 
characterized by the tufted indumentum of trichomes with a single adjoining 
basal cell and the unisexual florets, mostly in different specimens (dioecy). Five 
subgenera are currently recognized: Baccharis (180 species), Molina (Pers.) 
Heering (130 species), Pteronioides Heering (60 species), Stephananthus (Lehm.) 
Heering (four species), and Tarchonanthoides Heering (23 species). A taxonomic 
revision and a phylogenetic analysis of Baccharis subgen. Tarchonanthoides 
are underway. This work summarizes the first results of the taxonomic revision. 
Baccharis subgen. Tarchonanthoides is characterized by the corollas of female 
florets with five papillose teeth and by the nearly fully cleft style apex of male 
florets; moreover this subgenus lacks the tufted indumentum characteristic of 
most Baccharis species. The subgenus occurs in the South American grasslands 
and savannahs from Southeastern Brazil to Western Bolivia, south to Central and 
Eastern Argentina, with the greatest diversity in Southeastern Brazil and Uruguay. 
Currently, 23 species and 10 synonyms are recognized within the subgenus, as 
a result of the revision of the 33 names previously published, the description 
of three new species and the proposition of a new combination and status. 
Additionally seven lectotypes are chosen. Morphologically, the species of the 
subgenus are classified into four sections: Canescentes Giuliano (nine species), 
Coridifoliae Giuliano (nine species), Curitybensis Giuliano (three species), and 
Tarchonanthoides (Heering) Cuatrec. (two species). However, changes in the 
circumscription and composition of the sections may take place when results of 
the forthcoming molecular phylogenetic studies will be completed. 



34 Comp.Newsl. 50,2012 

Within-population variation of floral morphology in 
Aster hispidus var. tubulosus: its relationship to microhabitat 

and to pollinators 

Sayaka Nakagawa & M. Ito 



Department of General Systems Studies, 

Graduate School of Arts and Sciences, The University of Tokyo, 

3-8-1, Komaba, Meguro-ku, Tokyo 153-8902, Japan 

hayai-kame@hotmail.co.jp 

Within-population variation in floral morphology provides an ideal study system 
to understand plant diversification through adaptive evolution. Aster hispidus var. 
tubulosus, an endemic variety of A. hispidus complex in Japan, shows extreme 
within-population variation in floral morphology, from ligulate to long tubular 
ray florets in the outermost wheel of the capitulum. In this study, we examined 
whether this variation can be attributed to environmental factors of their habitats. 
The study population was subdivided into small patches alongside the river, which 
differed in the micro-environment and the frequency of floral types. According to 
a two-year survey, we found that the frequency of individuals with long tubular 
ray florets is negatively correlated with the degree of coverage of the patches. 
We hypothesized that the observed correlation was due to different selection 
pressures acting on floral morphology (presumably genetically controlled), which 
might depend on the degree of coverage of the patches. We also investigated 
whether morphology of outermost florets affects pollinator attraction in the field. 
As a result, we found that the ligulate ray florets have an advantage in attracting 
pollinators over the long tubular ray florets. We will discuss possible adaptive 
significance of long tubular ray florets and a mechanism of maintaining continuous 
floral variation in this species. 



Comp. Newsl. 50, 2012 35 

Phylogenetic study of Mikania Willd. 

(Compositae - Eupatorieae): 

a preliminary analysis 

Caetano Troncoso Oliveira, Benoit Loeuille & Jose Rubens Pirani 



Laboratorio de Sistematica Vegetal, Departamento de Botanica, 

Instituto de Biociencias, Universidade de Sao Paulo, Rua do Matao Travessa 14, 

321, 05508-090, Sao Paulo, SP, Brazil 

caetano.to@gmail.com 

Mikania Willd. contains ca. 450 species, it is the largest genus of Eupatorieae 
and one of the greatest of Compositae. It has a pantropical distribution, but its 
richness is concentrated in South America. The species of the genus are vines or 
shrubs, with heads composed of four florets and four phyllaries. Because of its 
size, taxonomic revisions are difficult to be performed. Infrageneric classifications 
have been proposed since the nineteenth century, mainly based on species habits, 
leaf shape and arrangement of the heads. Although there are few doubts about 
the monophyly of the genus, phylogenetic studies have never been performed to 
evaluate whether suggested infrageneric taxa are monophyletic. The present work 
is a preliminary analysis towards a more comprehensive molecular phylogeny 
of the genus. Sequences of approximately 900 bp of the Internal Transcribed 
Spacer (ITS) were obtained for 20 species of Mikania. The parsimony analysis 
resulted in four cladograms in which two mains clades with high jackknife 
support emerged, each one comprising species of different habits and types of 
inflorescence. However, it is necessary to include more terminals and to analyze 
additional molecular markers in order to have a clearer picture of the relationships 
within Mikania. 



36 Comp. Newsl. 50, 2012 

On the identity of Pluchea incisa Elmer, with 

notes on the genera Blumea and Pluchea 

(Asteraceae-Inuleae) 

Arne A. Anderberg 

Department of Phanerogamic Botany 

Swedish Museum of Natural History 

P.O. Box 50007, SE-104 05 Stockholm, Sweden 

arne.anderberg@nrm.se 

Abstract 

The species Pluchea incisa Elmer is discussed after investigation of floral micro- 
characters of the stamens and style. It is concluded that it is not a member of 
Pluchea or the Inuleae-Plucheinae, but instead belongs in the genus Blumea of 
the Inuleae-Inulinae. 

Introduction 

The species Pluchea incisa was described by Elmer (1908) as belonging to 
Candolle's Pluchea section Hebephora (Candolle 1836), a small group with 
two species {P. hirsuta and P. scabridd). The notion that Blumea and Pluchea 
are two closely related genera of the Inuleae is a traditional view founded on 
the fact that in both, the outer female florets are tubular or filiform, whereas the 
inner ones are male or bisexual. To Merrill & Merritt (1910) Pluchea incisa 
instead appeared to be a species of Blumea related to B. chinensis (L.) DC, and it 
was moved to that genus as B. incisa (Elmer) Merr. No characters in particular 
were put forward in support of this opinion, other than an impression of overall 
similarity. In a revision of Pluchea, King- Jones (2001) mentioned that Elmer's 
Pluchea incisa was likely belonging to another genus but without giving any 
statement of an alternative placement. Among other things she put forward the 
yellow flowers, subscandent growth habit and capitula arranged in a racemose 
fashion as characters indicating a placement outside of Pluchea. In this context it 
may be worthwhile to recapitulate the difference between the two genera Blumea 
and Pluchea. 



Comp.Newsl. 50,2012 37 



Taxonomic background 

In the first cladistic study of the Inuleae, Anderberg ( 1 989) concluded that Blumea 
and Phichea belonged to different major lineages. The analyses were based on 
morphological characters, several of which were floral micro-characters. Among 
other things, the styles of Pluchea were found to have sweeping-hairs that were 
rather long and distinctly obtuse, and also distributed a long distance down the 
shaft below the bifurcation of the style. In Blumea, the well-developed sweeping- 
hairs were more or less acute and confined to the style branches or sometimes 
diminishing in size and extending down to the bifurcation or just below. Acute 
sweeping-hairs like in the Blumea kind of styles were found in genera of the 
subtribe Inulinae in e.g. Inula and Pulicaria, whereas the Pluchea kind of style 
was typical of genera such as Epaltes, Laggera, Nicolasia, and Sphaeranthus . The 
stamen filament collars in many Pluchea species have characteristically swollen 
cells, whereas the filament collars in stamens of Blumea are always of a flat, 
quadrangular shape. An interesting discovery was that the Pluchea kind of styles 
was also found in two African species of Blumea (B. cafra and B. bovei), and the 
genus Doellia was later described to accommodate these two species (Anderberg 
1995). In some modern African flora treatments, Doellia has not been accepted 
and its species are still included in Blumea (Beentje 2002, 2006), disregarding the 
difference in style morphology. The conclusion that Blumea and Pluchea belonged 
to different evolutionary lineages, and that Doellia belonged in the Plucheinae 
clade together with Pluchea rather than with Blumea, has been corroborated by 
analyses of DNA sequence data by Anderberg et al. (2005) in which two major 
clades of genera were identified, corresponding to the subtribes Inuleae-Inulinae 
and Inuleae-Plucheinae, respectively. All genera of the Inulinae clade also showed 
a 3 bp insertion in ndhF that was not present in any genus of the Plucheinae clade. 
Interestingly, Doellia with its plucheoid stylar sweeping-hairs (Anderberg 1995, 
Fig. 1 F) also lacked this 3 bp DNA insertion. The distinction between Blumea and 
Pluchea is clear-cut, and if DNA sequences of ndhF are not obtainable, the shape 
of the stylar sweeping-hairs and filament collar cells may give a good indication 
of which genus a plant belongs to. 

A first detailed investigation of styles in loose flowers from the type specimen of 
Pluchea incisa could either give support to Merrill's opinion, or indicate that the 
plant belongs to another genus as suggested by King-Jones (2001). 

Methods, Results & Discussion 

Flowers were soaked in water, dissected and mounted in Hoyer's solution 
(Anderson 1954) for microscope investigation of floral micro-characters. The 



38 Comp. Newsl. 50, 2012 

anthers have almost unbranched tails, the endothecial tissue has radial wall 
thickenings and the filament collar is well developed and formed by slightly 
elongated quadrangular cells, and they are not swollen. The stylar sweeping-hairs 
are acute at the distal and median part of the style-branches but become shorter 
and more obtuse towards and just below the bifurcation where they disappear. 
Judging from the overall morphology of the plant, and by investigation of the 
floral micro-characters, I feel convinced that Merrill & Merritt (1910) were 
right in assuming that Pluchea incisa is a species belonging in Blumea. 

Investigated material 

Philippines. Benguet, Island Luzon, Baguio. Elmer 8396 (K, L). 

Acknowledgements 

I am grateful to the curators of the Kew (K) and Leiden (L) herbaria for sending 
material on loan. 

References 

Anderberg. A. A. 1989. Phylogeny and reclassification of the tribe Inuleae 
(Asteraceae). Can. J. Bot. 67: 2277-2296. 

Anderberg. A. A. 1995. Doellia, an overlooked genus in the Asteraceae- 
Plucheeae. Willdenowia 25: 19-24. 

Anderberg. A. A., Eldenas, P., Bayer, R. J. & M. Englund 2005. Evolutionary 
relationships in the Asteraceae tribe Inuleae (incl. Plucheeae) evidenced by 
DNA sequences of ndhF; with notes on the systematic position of some 
aberrant genera. Org. Div. & Evol. 5: 135-146. 

Anderson, L. E. 1954. Hoyer's solution as a rapid mounting medium for 

bryophytes. Bryologist 57: 242-247. 

Beentje, H. J. 2002. Blumea. In: Beentje, H. J. (ed.), Flora of Tropical East 
Africa, 319-322. Balkema, Rotterdam. 

Beentje, H. J. 2006. Blumea. In: Thulin, M. (ed.), Flora of Somalia 3: 495^196. 
Royal Botanic Gardens, Kew. 

Candolle, A. P. de 1836. Prodromus systematis naturalis regni vegetabilis 5. 
Treuttel & Wiirtz, Paris. 



Comp. Newsl. 50, 2012 39 



Elmer, A. D. E. 1908. A century of new plants. Leaflets Philippine Bot. 1(16): 

272-374. 

King-Jones, S. 2001. Revision of Pluchea Cass. (Compositae, Plucheeae) in the 
Old World. Englera 23: 1-136. 

Merrill, E. D. & M. L. Merritt 1910. The flora of Mount Pulog (concluded). 
Philippine J. Sci. 5: 371^107. 



40 Comp. Newsl. 50, 2012 

Ray-florets in Chiliadenus (Asteraceae- 

Inuleae), discovered. 

An epigenetic phenomenon? 

Arise A. Anderberg 



Department of Phanerogamic Botany 

Swedish Museum of Natural History, P. O. Box 50007 

SE-104 05 Stockholm, Sweden 

arne.anderberg@nrm.se 



Abstract 

Two specimens of Chiliadenus rupestris (Asteraceae, Inuleae) having capitula 
with one or two ray-florets have been discovered. The genus Chiliadenus is 
one of few in the Inuleae-Inulinae where capitula are consistently consisting of 
hermaphroditic disc-florets only. A discussion of the trait is presented, and the 
morphology and characteristics of the ray-florets are described and illustrated. 

Introduction 

The species of Chiliadenus Cass, have formerly been included in either Jasonia 
Cass, or Varthemia DC, but the genus was again recognized as distinct from these 
two by Brullo (1979), who e.g. stated that one difference between Jasonia and 
Chiliadenus was that the former has radiate heads with distinct ray-florets whereas 
the latter has discoid heads with hermaphroditic tubular disc-florets only. The 
genus Chiliadenus has nine species distributed around the Mediterranean, and has 
consistently (e.g., Quezel & Santa 1963, Tutin 1976, Feinbrun-Dothan 1978, 
Alavi 1983, Bolos & Vigo 1995, Boulos & Hind 2002, Vogt 2002, Anderberg 
& Eldenas 2007, Blanca 2009) been presented as a genus with discoid capitula 
composed of hermaphroditic tubular disc-florets only. In taxonomic treatments 
and determination keys, there is no information on variation in this character and 
to my knowledge there is no reference in the literature mentioning presence of 
ray-florets in species today placed in Chiliadenus. Two specimens of Chiliadenus 
rupestris (Pomel) Brullo have now been found, both with capitula provided with 
ray-florets. 



Comp. Newsl. 50, 2012 



41 



Material and methods 

The two radiate specimens were both collected at Beni-Snassen in Northern 
Morocco [Jury 13031 (BC, RNG, S) and Jury 15481 (RNG)]. Ray-florets were 
soaked in water and mounted in Hoyer ' s solution (Anderson 1 954) for microscope 
investigation. 

Results 



In the cladistics analysis of the Inuleae of Anderberg (1991), which was based 
on morphological data, three characters relating to heterogamous capitula and 
their marginal florets were coded as "unknown'' in Chiliadenus, i.e. the shape of 
marginal florets, the sex of marginal florets, and the shape of ray-floret epidermis 
cells. In the light of the new discovery additional and more complete data can be 
provided. 




Fig. 1. Ray-floret from Chiliadenus rupestris. Voucher: Jury 13031 (RNG). 



42 Comp. Newsl. 50, 2012 



Description of ray-florets. Ray-florets when present (Fig. 1), 1-2 per capitulum, 
ca. 10 mm long, pistillate; lamina short, elliptic, 5x2 mm, in-rolled (at least 
on herbarium specimens), 4-5-veined, apically with three or four short lobes; 
epidermis cells elongated in outline, with surface striations and minute marginal 
denticulations, without linear crystals, not crested; tube ca. 4 mm. Style bifid; 
style-branches elongated, flattened, somewhat wider distally, adaxially with two 
distally confluent bands of stigmatic tissue. Pappus pale yellowish-red, consisting 
of several barbellate bristles in one row surrounded by a series of free, narrowly 
triangular scales. Cypselas (immature) 2 mm long, ellipsoid, glandular-hairy, 
proximally with elongated twin-hairs. Carpopodium small. 

Discussion 

Not all capitula on the radiate specimens of Chiliadenus rupestris seem to have 
them, and in those capitula where there are rays they are only one or two per 
capitulum. The ray-florets have a distinct lamina about half as long as the ray- 
floret (lamina in-rolled on the dried herbarium specimens) and well-developed 
styles. 

The largest analysis of molecular data from the Inuleae-Inulinae is that of ndh¥ 
sequences from 160 species by Englund et al. (2009). In this study, Chiliadenus 
was found to be the sister-group of Dittrichia, and the two genera were part of a 
larger clade comprising also Jasonia Cass, and Pulicaria Gaertn. These genera 
have capitula with ray-florets, but in Jasonia they are neuter without any style. 
Apart from Jasonia, a few other genera of the Inuleae also have ray florets without 
a style, i.e. Anvillea DC, Perralderia Coss., and Iphiona Cass., but these all 
belong to different monophyletic lineages and apparently the style has been 
lost in ray-florets on at least three different occasions. As concluded by Brullo 
( 1 979), Jasonia and Chiliadenus are not congeners, and it is also clear from DNA 
analysis that they do not form a monophyletic group, as the latter is more closely 
related to Dittrichia Greuter than to Jasonia. Dittrichia rays are pistillate, like 
the Chiliadenus specimens found here. Discoid capitula and loss of ray-florets 
probably evolved in the ancestor of Chiliadenus after it had differentiated 
from the ancestor of Dittrichia. The apomorphic trait was then inherited by the 
offspring as they evolved into the nine discoid species of today. Englund et al. 
(2009) included five of the nine species of Chiliadenus, and found a trichotomy 
with the Algerian C. hesperius (Maire & Wilczek) Brullo and the Spanish C. 
saxatilis (Lam.) Brullo unresolved in relation to a clade formed by the Maltese 
C. bocconei Brullo, the Libyan-Egyptian C. candicans (Delile) Brullo, and the 
Moroccan C. rupestris. Therefore it can be concluded that the radiate specimens 
of C. rupestris are not representing an archaic part of the genus retaining its rays 



Comp.Newsl. 50,2012 43 



as a symplesiomorphy, but they belong to a more derived group of taxa and the 
rays have apparently reappeared secondarily. 

Normally radiate species may have individuals, populations or subspecific taxa 
without rays, e.g. Anvillea garcinii DC. (Anderberg 1982), Senecio vulgaris L. 
(Chater & Walters 1 976). Rays may be lost due to adaptation to arid environments 
like in some south Algerian representatives of Anvillea garcinii ssp. radiata 
(Coss. & Dur.) Anderb., or species may be polymorphic for other reasons. In 
Senecio vulgaris it has been shown that the radiate condition is dominant over the 
discoid, and that a change from homozygous for dominant alleles or heterozygous 
for dominant alleles to homozygous for recessive alleles may result in the loss 
of rays. Two gene loci (RAY1, RAY2) and a number of modifier genes seem 
to be active in the presence /absence of rays in Senecio vulgaris (Gillies et al. 
2002). Homozygotic dominant alleles give long rays, heterozygotic dominant 
give short rays, and homozygotic recessive results in loss of ray-florets. There 
seems to be one or two genes regulating the formation of ray-florets also in other 
Compositae, such as Gerbera L. (Laitinen et al. 2006) and Layia Hook. & Arn. 
(Ford & Gottlieb 1990), but the absence of rays in a plant does not mean it has 
lost the necessary regulatory genes (Baldwin 2005). Gillies et al. (2002) tested 
the hypothesis that CYCLOIDEA (CYC) gene homologues were involved in the 
formation of ray-florets in Senecio. This gene family is known to be involved in 
formation of the capitulum, and according to Carlson et al. (20 1 1 ) there have been 
several rounds of duplication of CYC-like genes in the Asteraceae, to some extent 
involved in the formation of zygomorphic flowers, with radiate species having 
more copies than discoid species. Chapman et al. (2012) presented evidence for 
several paralogues of CYC2-like genes being present in the Asteraceae. 

Situations where genera with only discoid species where some suddenly display 
individuals with ray-florets are apparently very unusual and I have not been able 
to find any discussion of this phenomenon in the literature. Nothing is known 
about the genetic mechanisms at work specifically in Chiliadenus, but it is 
reasonable to believe that it follows the same general patterns as described for 
other Asteraceae. If it conforms to Senecio vulgaris, where also the heterozygotic 
condition for dominant alleles result in presence of rays, one would expect the 
ancestor of Chiliadenus to be homozygotic recessive as no rays are formed. If so, 
there would not be any dominant ray-floret alleles present, and the mechanisms 
behind the occurrence of ray-florets in C. rupestris would have to be explained 
in some other way. One bold and perhaps far-fetched hypothesis could be that all 
Chiliadenus have the ray-floret genes, but that epigenetic suppression of these 
regulator or associated modifier genes has inhibited ray-floret development until 
some circumstance made ray-florets reappear in the Beni-Snassen population. 
Apparently this phenomenon may have many different causes and the mechanisms 



44 Comp. Newsl. 50, 2012 



behind them are far from clear. Perhaps these plants may one day be used for 
genetic research to shed more light on the issue on why ray-floret phenotypes are 
formed. 



Acknowledgements 

I am grateful to Emma Hulten for the illustration of the ray-florets. I also want to 
acknowledge Paula Elomaa, Helsinki, Richard Abbott, St. Andrews, and Mark 
Chapman, Oxford, for inspiring communications. 

References 

Alavi, S. A. 1983. Asteraceae. Pp. 1^55 in: Jafri, S. M. H. & A. El-Gadi (eds.), 
Flora of Libya 107. Al Faateh University, Tripoli. 

Anderberg, A. A. 1991. Taxonomy and phylogeny of the tribe Inuleae 
(Asteraceae). PI. Syst. Evol. 176: 75-123. 

Anderberg, A. A. & P. Eldenas 2007. Inuleae. Pp. 374-391 in: Kadereit, J. W. 
& C. Jeffrey (eds.), The families and genera of vascular plants 8. Springer- 
Verlag. Berlin Heidelberg New York. 

Anderberg, A. A. 1982. The genus Anvillea (Compositae). Nord. J. Bot. 2: 
297-305. 

Anderson, L. E. 1954. H oyer's solution as a rapid mounting medium for 

bryophytes. Biyologist 57: 242-247. 

Baldwin, B. 2005. Origin of the serpentine-endemic herb Layia discoidea from 
the widespread/,, glandulosa (Compositae). Evolution 59: 2473-2479. 

Blanca, G. 2009. Chiliadenus. Pp. 380-381 in: Blanca, G., Cabezudo, B., Cueto, 
M., Fernandez Lopez, C. & C. Morales Torres (eds.), Flora vascular 
de Andalucia Oriental 4. Verbenaceae-Asteraceae. Consejeria de Medio 
Ambiente, Junta de Andalucia, Sevilla. 

Bolos, O. de & J. Vigo 1995. Flora dels Pa'isos Catalans 3. Editorial Barcino, 
Barcelona. 

Boulos, L. & D. J. N. Hind 2002. Compositae. Pp. 134-320 in: Boulos, L. (ed.), 
Flora of Egypt 3. (Verbenaceae-Compositae) . Al Hadara Publishing, Cairo. 

Brullo, S. 1979. Taxonomic and nomenclatural notes on the genera Jasonia 
Cass, and Chiliadenus Cass. (Compositae). Webbia 34: 289-308. 

Carlson, S. E., Howarth, D. G. & M. J. Donoghue 2011. Diversification of 
CYCLOIDEA-like genes in Dipsacaceae (Dipsacales): implications for 



Comp.Newsl. 50,2012 45 



the evolution of capitulum inflorescences. BMC Evol. Biol. 2011 11: 325. 
doi:10.1186/1471-2148-ll-325. 

Chapman, M. A., Tang, S.-X., Draeger, D., Nambeesan, S., Shaffer, H., 
Barb, J. G., Knapp, S. J. & J. M. Burke 2012. Genetic Analysis of Floral 
Symmetry in Van Gogh's Sunflowers Reveals Independent Recruitment 
of CYCLOIDEA Genes in the Asteraceae. PLoS Genet. 8(3) el 002628. 
doi: 10. 1371/journal.pgen. 1002628 

Chater, A. O. & S. M. Walters 1976. Senecio. Pp. 191-205 in: Tutin, T. G, 

Heywood, V. H., Burges, N. A., Moore, D. M., Valentine, D. H., Walters, 
S. M. & D. A. Webb (eds.), Flora Europaea 4. Plantaginaceae to Compositae 
(and Rubiaceae) . Cambridge University Press, Cambridge. 

Englund, M., Pornpongrungrueng, P., Gustafsson, M. H. G. & A. A. 

Ainderberg 2009. Phylogenetic relationships and generic delimitation in 
Inuleae subtribe Inulinae (Asteraceae) based on ITS and cpDNA sequence 
data. Cladistics 25: 319-352. 

Feinbrun-Dothan, N. 1978. Flora Palaestina 3. Ericaceae to Orchidaceae. The 
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Ford, V. S. & L. D. Gottlieb 1990. Genetic studies of floral evolution in Layia. 
Heredity 64: 29-44. 

Gillies, A. C. M., Cubas, P., Coen, E. S. & R. J. Abbott 2002. Making rays 
in the Asteraceae: genetics and evolution of radiate versus discoid flower 
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(eds.), Developmental genetics and plant evolution. Taylor & Francis, 
London. 

Laitinen, R. A. E., Broholm, S., Albert, V. A., Teeri, T. H. & P. Elomaa 

2006. Patterns of MADS4}ox gene expression mark flower-type 
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6:11. doi: 10. 11 86/1471-2229-6-11. 

Quezel, P. & S. Santa 1963. Nouvellejiore del'Algerie etdes regions desertiques 
meridionales. Editions du Centre National de la Recherche Scientifique, 
Paris. 

Tutin, T. G. 1976. Jasonia. P. 138 in: Tutin, T G, Heywood, V H., Burges, 
N. A., Moore, D. M., Valentine, D. H., Walters, S. M. & D. A. Webb 
(eds.), Flora Europaea 4. Plantaginaceae to Compositae (and Rubiaceae) . 
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Vogt, R. 2002. Chiliadenus. Pp. 637-638 + key in: Valdes, B., Rejdali, M, 
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46 Comp. Newsl. 50, 2012 

The genus Cavea, an addition to the tribe 

Gymnarrheneae 

(Asteraceae-Gymnarrhenoideae) 

Arne A. Anderberg 1 & Jan I. Ohlson 2 

'Department of Phanerogamic Botany 

department of Molecular Systematics 

Swedish Museum of Natural History 

P.O. Box 50007, SE-104 05 Stockholm, Sweden 

arne.anderberg@nrm.se 

Abstract 

The systematic position of the yet unplaced Cavea tanguensis was investigated 
by analysis of DNA sequences from the plastid gene ndhF. It is concluded that 
Cavea tanguensis is the sister of Gymnarrhena micrantha, and that the genus 
Cavea, which has been notoriously difficult to place with any certainty, is a second 
member of tribe Gymnarrheneae (Asteraceae-Gymnarrhenoideae). 

Introduction 

The present paper is one in a series of studies aiming at finding the systematic 
position for aberrant and odd representatives of the Asteraceae, such as Caesulia 
Roxb. (Anderberg et al. 2005), Dipterocome Fisch. & C. A. Mey. (Anderberg et 
al. 2007), Feddea Urb. (Cariaga et al. 2008), Gymnarrhena Desf. (Anderberg 
et al. 2005), Nanothamnns Thomson (Anderberg & Pandey 2008), Psednotrichia 
Hiern (Anderberg & Karis 1995), and Rhodogeron Griseb. and Sachsia Griseb. 
(Hong et al. 2004). 

One remaining systematic enigma in the family is the monotypic genus Cavea W. 
W. Sm. & J. Small from the Himalayas (Fig. 1). It is one of the last rogue genera 
of the Asteraceae, i.e. genera for which the systematic position is unclear and yet 
unsolved and not tested in a phylogenetic framework based on DNA sequence 
analyses. In some more recent treatments of the family (Jeffrey 2009, Pelser 
& Watson 2009, Chen & Anderberg 2011) Cavea has been treated as a genus 
incertae sedis, either belonging to the Cichorioideae or the Asteroideae, and its 
tribal relationship has also earlier been much discussed. Most authors have been 
eager to exclude it from the group they have been studying, but none willing to 



Comp. Newsl. 50, 2012 47 



accept it. 

The morphological features of Cavea have made it difficult to place in relation to 
other genera. Bremer ( 1 994), who listed Cavea among the Asteroideae unassigned 
to a tribe, noted that its filament collars were poorly developed and its endothecial 
tissue thickenings inconspicuous or absent like in the Cichorioideae tribes. He also 
stated that its style had stigmatic lines in two bands unlike the Cichorioideae, but 
typical of the Asteroideae. Jeffrey (2007) regarded Cavea as most likely related 
to the Inuleae in spite of its Carduoid facies, and later (Jeffrey 2009) pointed out 
its breeding system as being unique with both dioecious and monoecious plants 
in the same population. 

In spite of all the detailed morphological investigations, the systematic position 
of Cavea has remained unsolved. Therefore it was the more intriguing to try and 
shed light on these issues by analysis of DNA sequence data. The results were 
surprising. 

Taxonomic history 

The species Saussurea tanguensis J. R. Drumm. was described in 1910 and placed 
in Cynaroideae-Carduineae (Drummond 1910). Later the new genus Cavea was 
described based on Drummond's species (Smith & Small 1917), and with great 
hesitation placed near Pluchea of the Inuleae. The authors discussed the tendency 
in Cavea towards dioecy (large amount of capitula with no male florets), and that 
the Inuleae in their pollen presentation mechanism were closer to the Cynareae 
and Mutisieae, and also that the style and stamens would place Cavea nearer to 
Gochnatineae of the Mutisieae, than to Saussurea. 

Ling & Chen (1965) agreed with Smith & Small in that the genus belonged in the 
Inuleae-Plucheinae but provided an amplified and much more detailed description 
of the genus. Among other things, they noted that the plants showed different 
breeding systems in the same population and that some were smaller in size, 
sometimes even stemless (Ling & Chen 1965, pi. 19 fig. 14). 

Dittrich (1977) as well did not accept Cavea in the Cardueae and instead 
suggested it to have an uncertain position in the Inuleae. This view was not 
shared by Merxmuller et al. (1977) who excluded it from the Inuleae because 
of aberrant pollen morphology with infratectal bacules ["infrategillar bacules"] 
(Merxmuller et al. 1977, p. 579). Based on his thorough experience of Inuleae 
pollen ultrastructure, Leins (1971) concluded that Cavea pollen was sufficiently 
different from what he had seen in the Inuleae to merit its exclusion from that 
tribe. Anderberg (1991) also kept Cavea out of the Inuleae s.str., and from his 
Plucheeae. This opinion was based on the presence in Cavea of a deeply lobed 



48 Comp.Newsl. 50,2012 



corolla, undifferentiated anther filament collar, indistinct or missing endothecial 
thickenings, and also supported by the infratectal pollen wall (see above). 

Material and methods 

DNA was extracted from leaves taken from herbarium specimens of Cavea 
tanguensis [Vouchers: Rock 16868 (S); Sinclair & Long 5423 (E); Ludlow, 
Sherriff & Hicks 16309 (E); Ludlow, Sherriff & Hicks 20793 (E)]. To test the 
tribal position of Cavea tanguensis in the family, four obtained ndhF sequences 
were analyzed together with the data set (184 taxa) from Anderberg et al. (2005) 
and with an additional number of unpublished sequences, mainly from the Inuleae, 
a total of 258 sequences of taxa representing all Asteraceae tribes. 

Molecular methods. DNA extraction was carried out with the QIAGEN DNeasy 
Plant Mini Kit using the manufacturer's protocol. PCR reactions were performed 
with PuReTaq Ready-To-Go PCR Beads. Amplification was performed under the 
following settings: 95°C 5 min, followed by 4 cycles of 95°C 30 sec, 54°C 30 sec, 
72°C 1 min 15 sec; 4 cycles of 95°C 30 sec, 52°C 30 sec, 72°C 1 min 15 sec and 36 
cycles of 95°C 30 sec, 49°C 30 sec, 72°C 1 min 15 sec, with a final extension step 
at 72° 8 min. Purification of PCR products was done with the ExoFast enzymatic 
purification kit (Fermentas Life Sciences) following the manufacturer' s protocol. 
Sequencing reactions were made with the same primers as in the amplification, 
using the BigDye Terminator v.3.1 Cycle Sequencing Kit. Unincorporated dye 
terminators were removed using QIAGEN's DyeEx 96 Kit. Fragments were 
separated and analyzed on an ABI 3130x1 Genetic Analyzer. Primers used for 
PCR and sequencing of ndhF are presented in Table 1 . The four new sequences 
have been submitted to GenBank (Accession numbers JQ922540-JQ922543). 

Alignment. Alignment of ndhF was performed with the BioEdit software (Hall 
1999) ver. 6.0.5. The aligned ndhF data set included 258 sequences with several 
representatives of each tribe of the Asteraceae. 

Phylogenetic analyses. The alignment of the 258 ndhF sequences resulted in a 
data matrix with 2289 sites which was analyzed with parsimony jackknifing using 
the software XAC (Farrjs 1997) with the following settings: 1000 replications, 
each with branch-swapping and 10 random-additions of sequences. For the 
analysis, Boopis (Calyceraceae) was used as outgroup (F arris 1972). 

Results 

The results of the parsimony jackknife analysis (Fig. 2) showed that 703 of the 
2289 sites were informative, and that the four sequences of Cavea formed a 



Comp.Newsl. 50,2012 49 



monophyletic group (100 % jackknife support), with Gymnarrhena micrantha as 
their sister group. The position of the Gymnarrhena - Cavea clade was congruent 
with the position of the former in the Cichorioid tribal complex in Anderberg et 
al. (2005). 

Discussion 

The strongly supported close relationship between Cavea and Gymnarrhena was 
unexpected as the two have no obvious similarities. 

Gymnarrhena micrantha is a small rosulate, stemless herb with linear lanceolate 
entire leaves and two kinds of capitula. One kind being female, cleistogamous and 
subterranean, the other kind is situated in the soil surface and provided with both 
female and male florets. Funk & Fragman-Sapir (2009) reported that Gymnarrhena 
can survive quite harsh environmental changes and that its amphicarpy may be an 
autapomorphic adaptation to desert life. The genus has a distribution over large 
parts of North Africa and the Middle East. Gymnarrhena was once placed in the 
Inuleae (Bentham 1873, Hoffmann 1890), just like Cavea, but later authors have 
not accepted it in treatments of that tribe (e.g. Leins 1973, Merxmuller et al. 
1977, Anderberg 1991 ). The genus Gymnarrhena has been difficult to place, but 
DNA analysis has shown it to take a rather isolated position in the Cichorioideae 
(Anderberg et al. 2005). Earlier Funk & Panero (2002) had described a new tribe 
and subfamily (Gymnarrhenoideae-Gymnarrheneae) to accommodate this small 
monospecific evolutionary lineage, and placed it outside of the Cichorioideae- 
Corymbieae-Asteroideae. Gymnarrhena has an unusual aspect and its habit, gross 
morphology and its micromorphology have not provided any clues as to what its 
closest relative may be. The pollen was investigated by Wortley et al. (2007) 
in search of synapomorphies that could support its position in the Asteraceae 
phylogeny as evidenced by molecular data, but the study was not very conclusive 
as they found the ultrastructure not matching that of the Cichorioideae, but with 
some features approaching those in Cotymbium (Corymbieae) stated to be the 
sister group of the Asteroideae. 

The results of the present investigation placing the likewise enigmatic genus 
Cavea as sister to Gymnarrhena are of course interesting. Although they are 
both autapomorphic and without any obvious relatives as judged from their 
morphological characters, they must both have some taxon to which they are 
more related than to others. An explicit hypothesis of a close generic relationship 
between the two has never been presented before, and it must be noted that the 
presence in Cavea of two kinds of capitula with a tendency towards dioecism, and 
the presence of two growth forms, one with distinct stem and the other acaulescent 
are striking given a position as sister to the likewise acaulescent Gymnarrhena. 



50 Comp.Newsl. 50,2012 



The latter also has two kinds of capitula, one with only female florets, and the 
other with male and female florets mixed, which Funk & Fragman-Sapir (2009) 
suggested to be possible clusters formed by reduction of several capitula sitting 
close together. The pollen of Cavea tanguensis has yet to be investigated, but it 
would be some support for its position if it could be shown to have infrategillar 
baculae like Gymnarrhena micrantha. 

In biogeographical terms, the two genera are allopatric. Gymnarrhena grows in 
deserts from North Africa to the Middle East, and Cavea on alpine meadows and 
gravelly places beside streams and glaciers on high altitudes in the Himalayas much 
further to the East. Both plants inhabit environments that are known to promote 
adaptations and morphological specialization. This distribution is the remains of 
a once continuous range of their ancestor, and the morphological characteristics 
of each of the two genera the result of long isolation in an eastern and western 
distribution. The addition of Cavea to the tribe Gymnarrheneae contributes to the 
understanding of the origin of that clade. From being an isolated Saharo-Sindian 
taxon, the origin of the tribe now seems to be in Asia, in an area that later broke 
up and formed the present day distributions of the two genera. 

Conclusion 

The present analysis of ndh¥ sequence data indicates with strong support that the 
genus Cavea is sister to the Saharo-Sindian genus Gymnarrhena. Both genera 
have an unusual morphology and several unique features that have made them 
difficult to place. It is concluded that Cavea is not a member of the Asteroideae as 
previously assumed, but part of the Cichorioideae tribal complex, likely a second 
genus in the small tribe Gymnarrheneae, and thus yet another Asteraceae genus 
incertae sedis may have found a secure resting place. 



Investigated material of Cavea tanguensis 

Tibet, Melong Gompa, 2.VII.1939, Gould 2233 (K); Tibet, Rama to Melong 
Gompa, 1. VII. 1939, Gould 2326 (K); Eastern Himalaya, Nam La, 23.1.1924, 
Ktngdon Ward 5977 (K); East Himalaya, Chumoleri, 12.IX.1912, Lepcha 455 
(E); Bhutan, Lingshi Dzong, 25. V. 1949, Ludlow, Sherriff & Hicks 16309 (E); 
Bhutan, Gafoola, Upper Pho Chu, 7.VII.1949, Ludlow, Sherriff & Hicks 16763 
(E); Tibet, Cho La, North side, 4. VII. 1949, Ludlow, Sherriff & Hicks 20793 
(E). China, Szechuan, Mont Konka, Risonquemba, Konkaling, V-VIII 1928, Rock 
16868 (E, S); Bhutan, Upper Mo Chu District, valley SW of Lingshi Dzong, 
Sinclair & Long 5423 (E, K); Tibet, Yanthang, 16.VII.1906, White s.n. (K). 



Comp. Newsl. 50, 2012 51 



Acknowledgements 

We are grateful to the Edinburgh and Kew herbaria for sending material on loan, 
and also to Ulf Swenson for drawing the tree. 

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Anderberg, A. A., Eldenas, P., Bayer, R. J. & M. Englund 2005. Evolutionary 
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Anderberg, A. A., Ghahremaninejad, F. & M. Kallersjo 2007. The enigmatic 
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Cariaga, K. A., Pruski, J. F., Oviedo, R., Anderberg, A. A., Lewis, C. E. & 
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Table 1. 

Primer sequences (5'-3') for ndhF. 

F = forward, R = reverse. 

Primer sequence 

AGG TAA GAT CCG GTG AAT CGG AAA C 

GTC TC A ATT GGG TTA TAT GAT G 

GGA GCT ACT TTA GCT CTT G 

GTT AAA CCT CCC ATA AGC ACC ATA TTC TGA C 

TAT GAT CCA ACC CTT TMT TTM TAT TCC 

CAT AGT ATT ATC TGA TTC ATA AGG ATA 

CCT ACT CCA TTT GGA ATT CCA TC 

All primers except were published in Kallersjo et al. (2000), except for primers 
520 and 1 750 that were published by Anderberg & Swenson (2003). Primers RJ1 
and RJ14 were designed by Ki-Joong Kim and Robert Jansen. 



Name 


Direction 


RJ1 


F 


5 


F 


5B 


F 


16 


R 


1650-Ast 


F 


1700 


R 


10B 


R 



54 



Comp.Newsl. 50,2012 



V 









Hert).No.&03-icso -« — 



Fig. 1. Cavea tanguensis. Scanned herbarium specimen. - Rock 16868 (S). 



Comp. Newsl. 50, 2012 



55 



Boopis 

Barnadesia 

Chuquiraga 

Dasyphyllum 

Doniophyton 

Schlechtendalia 

Adenocaulon 

Gochnatia 

Mutisia 

Onoseris 

Stifftia 

Leibnitzia 

Gerbera 

Piloselloides 

Nassauvia 

Trixis 

Perez/a 

Ainsliaea 

Echinops 

Tarchonanthus 

At racty lodes 

Carlina 

Saussurea 

Dipterocome 

Synurus 

Cirsium 

Cirsium 

Carthamus 

Centaurea 



Gymnarrhena 
Cavea R 16868 
Cavea LSH16309 
Cavea LSH20793 
Cavea SL5423 



LACTUCEAE (3) 
ARCTOTEAE (4) 
VERNONIEAE (8) 
SENECIONEAE (4) 
CALENDULEAE (3) 
GNAPHALIEAE (4) 
ANTHEMIDEAE (4) 
ASTEREAE (9) 
INULEAE (158) 
ATHROISMEAE (4) 
FEDDEAE (1) 
HELIANTHEAE slat. (22) 



Fig. 2. 



Parsimony jackknife tree based on ndhV sequence data showing the position of 
Cavea among the Asteraceae tribes, as sister group of Gymnarrhena. Support 
values > 50 % are shown for each clade. 



56 Comp. Newsl. 50, 2012 

Caputia, a new genus to accommodate four 

succulent South African Senecioneae 

(Compositae) species 

Bertil Nordenstam 1 & Pieter B. Pelser 2 

'Department of Phanerogamic Botany 

Swedish Museum of Natural History, Box 50007 

SE-10405 Stockholm, Sweden 

bertil.nordenstam@nrm.se 

2 University of Canterbury, School of Biological Sciences 

Private Bag 4800, Christchurch 8140, New Zealand 

pieter.pelser@canterbury.ac.nz 

Abstract 

The new genus Capittia B. Nord. & Pelser is described, with four species confined 
to South Africa. Its members are succulent perennial herbs or shrubs with more or 
less fleshy tomentose or glabrescent leaves. The genus has tussilaginoid as well 
as senecioid morphological characters, and takes strongly supported incongruent 
phylogenetic positions in nuclear and plastid phylogenies: sister to subtribe 
Brachyglottidinae or sister to the Synotoid group of Senecioninae, respectively. 
This suggests that the genus is potentially of hybrid origin. The four species are 
Capittia medley-woodii (Hutch.) B. Nord. & Pelser (type), C. tomentosa (Haw.) 
B. Nord. & Pelser, C. scaposa (DC.) B. Nord. & Pelser and C. pyramidata (DC.) 
B. Nord. & Pelser. 

Introduction 

One of the most interesting of the generic entities identified in our phylogenetic 
survey of the tribe Senecioneae (Pelser et al. 2007, 2010, Nordenstam et al. 
2009) is the Senecio medley-woodii group. There is no available generic name 
for this distinct assemblage of four succulent species, confined to South Africa, 
and some of which are quite spectacular and well known in cultivation (Rowley 
1994, Cullen et al. 2000, Eggli 2002). We here suggest the new generic name 
Caputia for this assemblage, an allusion to their geographical origin. The name has 
provisionally already been used, e.g. in Pelser et al. (2010). The old geographical 
name Caput bonae spei was often applied not only to the Cape of Good Hope 



Comp. Newsl. 50, 2012 



57 




Fig.l. Caputia medley-woodii. 

From Flow. PI. S. Afr. Plate 83 (1923), "Senecio Medley-Woodii" . 



58 Comp. Newsl. 50, 2012 



itself, but to the Cape Province or even the whole of South Africa. The generic 
name also alludes to Capelio, one of the early-diverging genera in the tribe, and 
herbaria with alphabetical generic order will then conveniently file these two 
genera close together. 

Discussion 

The members of Caputia used to be treated under Senecio L. (Harvey 1865, 
Hutchinson 1923, Rowley 1994, 2002) and one of them sometimes in Kleinia 
Mill. (De Candolle 1838, Harvey 1865, Marloth 1932). In phylogenies derived 
from nuclear as well as plastid DNA sequences the genus is only distantly related 
to Senecio s.str. In an ITS phylogeny of the tribe (Pelser et al. 2007), three Caputia 
species (included as Senecio medley-woodii, S. scaposus and S. pyramidatus) 
form a clade that is sister to subtribe Brachyglottidinae. This subtribe (which is 
not yet formally published) has a majority of genera in Australasia (Brachy glottis, 
Bedfordia, Dolichoglottis, Haastia, etc.), but also extensions into New Guinea 
(Papuacalia) and Chile (Acrisione). However, the plastid phylogeny presented 
by Pelser et al. (2007) indicates a close relationship between Caputia and the 
Asian genus Synotis in subtribe Senecioninae. This phylogenetic incongruence 
was later confirmed in a study that used a more extensive sampling of Senecioneae 
genera and characters (Pelser et al. 2010). The ITS/ETS phylogeny in the latter 
study places Caputia sister to the Brachyglottidinae ( 1 00% parsimony bootstrap 
support [BS]; 1.0 posterior probability [PP]). Again, plastid DNA sequence data 
suggest closer affinities with the Senecioninae and resolved Caputia sister to the 
Synotoid group (95% BS; 1.0 PP), which is composed of Austrosynotis (Tropical 
Africa), Cissampelopsis (Asia), Dauresia (Namibia), Humbertacalia plus a few 
related genera (Madagascar and Mascarenes), Mikaniopsis (Africa), and Synotis 
(E Asia especially Sino-Himalayan region). 

Pelser et al. (2010) suggested that ancient hybridization is the most likely 
explanation for the incongruent phylogenetic position of Caputia. Morphology 
provides some, although admittedly limited, support for this hypothesis. Subtribe 
Brachyglottidinae is characterized by 'tussilaginoid' features and among these 
are anthers with a cylindrical filament collar and polarized endothecial cell walls. 
In subtribe Senecioninae, however, mostly 'senecioid' features are found and 
these include balustriform filament collars and radial endothecial cell walls. The 
anthers of Caputia appear to be somewhat intermediate. Their filament collar is 
subcylindrical or weakly balustriform and the endothecium is transitional with 
radial and polarized thickenings. The disc-floret styles have continuous stigmatic 
areas, a feature characteristic of tussilaginoid genera and less common among the 
senecioid taxa. 






Comp.Newsl. 50,2012 59 



Taxonomy and Descriptions 

Caputia B. Nord. & Pelser, gen. nov. 

Erect branching shrubs, shrublets or thick-stemmed herbs. Leaves alternate, 
sessile, subcarnose to strongly succulent, flattened and thickish to terete, entire 
or coarsely dentate, mostly thinly to densely tomentose, sometimes glabrescent, 
sometimes persistently silvery white-felted. Capitula large, solitary or few to 
many in corymbose or thyrsoid synflorescences, radiate or discoid. Involucre 
cupshaped; phyllaries 8-13, uniseriate. Receptacle flat, glabrous, alveolate. Ray- 
florets absent or 8— 13(— 14), female, yellow; lamina strapshaped. Style bilobed. 
Cypsela ellipsoid-oblong, glabrous or with few scattered trichomes, sometimes 
more densely puberulous basally. Pappus bristles numerous, barbellate, white, 
persistent. Disc-florets numerous, hermaphroditic; corolla tubular, somewhat 
widening distally, 5-lobed; lobes triangular-ovate, midlined, apically subcucullate 
and somewhat papillate. Anthers basally obtuse to shortly sagittate; endothecium 
transitional with radial and polarized thickenings; filament collar subcylindrical 
or weakly balustriform; apical appendage ovate, flat. Style bifurcate with 
continuous stigmatic area inside the branches, apically subtruncate with few and 
short sweeping-hairs. Cypsela fusiform-ellipsoid, glabrous or puberulous. Pappus 
bristles numerous, white, persistent. Chromosome no. 2n = 20 (x = 10) with large 
chromosomes. 

Four species, distributed in mostly arid parts of Western and Eastern Cape 
Province, KwaZulu-Natal and Swaziland. 

Type species: Caputia medley-woodii (Hutch.) B. Nord. & Pelser. 

Key to the species of Caputia 

la. Stems and leaves with a dense silvery-white persistent tomentum. Capitula 
discoid 2.C. tomentosa 

lb. Plants araneose-tomentose, glabrescent. Capitula radiate 2 

2a. Leaves flattened, obovate — rhomboid 1. C. medley-woodii 

2b. Leaves terete (cylindrical, fusiform, sometimes apically flattened) 3 

3a. Stemless or short-stemmed shrublets. Capitula few (1 — 6) in racemose 
synflorescence 3. C. scaposa 

3b. Shrubs up to 180 cm high. Capitula numerous in conical or elongated 
synflorescence 4. C. pyramidata 



60 



Comp. Newsl. 50, 2012 




Fig. 2. Caputia tomentosa, flowering in National Botanic Gardens, Kirstenbosch, 
South Africa. Photo: B. Nordenstam 2007. 



Comp. Newsl. 50, 2012 



61 




Fig. 3. Caputia tomentosa. From Curtis Bot. Mag. 6063 (1873), 

"Senecio Haworthif '. 



62 Comp. Newsl. 50, 2012 



1. Caputia medley-woodii (Hutch.) B. Nord. & Pelser, comb. nov. 

Basionym: Senecio medley-woodii Hutch., Fl. PI. S. Afr. Plate 83 (1923). 

A branched shrub 0.5-2 m tall; stems thick, fleshy, often red or purplish, white- 
felted when young, glabrescent. Leaves sessile, obovate to rhomboid, 3-6 cm 
long, 1.5-3 cm wide, flattened but carnose, margins entire to coarsely dentate 
especially distally and somewhat undulate, apically mucronate-apiculate, basally 
somewhat cuneate, white-tomentose when young, becoming glabrous. Peduncle 
10-20 cm long with 1-4 capitula in corymbose arrangement, with small scattered 
bracts. Capitulum large, radiate. Involucre campanulate, with a few minute 
calyculus bracts; phyllaries 8-13, uniseriate, lanceolate-elliptic-ovate, 12-15 mm 
long, 3-6 mm wide, somewhat woolly, subcoriaceous, obtuse. Receptacle flat to 
somewhat convex, minutely alveolate with shortly fringed margins and a small 
central pit. Ray-florets ca. 12 (10-14), tube cylindric, 5-6 mm long, glabrous or 
with scattered short trichomes, lamina 1.5 cm long, bright yellow, 6-8-veined. 
Disc-florets 30-70, corolla 12 mm long, yellow or brownish-yellow, gradually 
widening upwards; lobes 1.5 mm long with a median resin duct. Style branches 
2 mm long, apically truncate and minutely penicillate. Cypsela 6-7 mm long, 
fusiform, glabrous, 5-ribbed. Pappus bristles 9-10 mm long, white, persistent. 
Chromosome no. 2n = 20 (Afzelius 1967). - Fig. 1. 

Selected collections (herbarium abbreviations as in Holmgren et al. 1990): 

SOUTH AFRICA, NATAL: /2930DA Pietermaritzburg/, Craiglea, opposite 
Monteseel to the E and Table Mountain to the W, edge of escarp, 19.VI.1977, P. C. 
V. de Toit 2437 (K); Oribi Flats, top of Oribi Gorge, IV. 1 937, A. P. D. McClean 563 
(K); Ngotshe, Lebombo Mts, Majezind area, 1 750 ft, 8. VII. 1 962, C. J. Ward 4 1 83 
(K); /3030 Port Shepstone/, granite hill 4 miles inland Umtentweni, 6.VIII. 1967, R. 
G. Strey 7617 (K, S); Natal, Port Shepstone, Fairacres Estate on Oribi Flats, krans 
overlooking deep Umzimkulwana Valley, 22. VII. 1953, R. A. Dyer 5416 (K); near 
Murchison, reed. X.1884, Medley-Wood 3065 (K); Krantz Kloof, 23.VIII.1915, 
J. M. Wood 13247 '," Medley- Woodii Hutchinson n.sp." (K 1 specimen, annotated 
by Hutchinson, syntype); Natal, Inanda, comm. VIII. 1879, J. M. Wood 555, 
"S. Medley-Woodii Hutch." (Hutchinson scripsit, K several specimens, syntypes). 

C. medley-woodii is near-endemic in KwaZulu-Natal, occurring also in 
neighbouring Eastern Cape and Swaziland. The habitats are granite and other 
rocky outcrops, river gorges etc. at lower altitudes up to 600 m.s.m. (Hilliard 
1977). 

Hutchinson (1923) named it for John Medley Wood (1827-1915), a Natal 
botanist of distinguished repute. The species is widely cultivated as a handsome 
succulent in greenhouses (Rowley 1994, 2002). 



Comp. Newsl. 50, 2012 



63 




Fig. 4. Caputia scaposa. From Curtis Bot. Mag. 401 1 (1 843), 
"Senecio calamifolius" . 



64 



Comp.Newsl. 50,2012 




Fig. 5. Caputia pyramidata. From Curtis Bot. Mag. 5396 (1863), 
"Senecio pyramidatus" '. 



Comp.Newsl. 50,2012 65 

2. Caputia tomentosa (Haw.) B. Nord. & Pelser, comb. nov. 

Basionym: Kleinia tomentosa Haw., Syn. PI. Succ: 314 (1812). 

Syn.: Cacalia tomentosa Haw. Misc. 189 (1803) non Jacq. 1775 (= Adenostyles 
alba) nee non Vill. 1779 nee non L.fil. 1782. 

Cacalia canescens Willd., Enum. Hort. Berol. Suppl. 427. (1814). 

Cacalia haworthii Sweet, Hort. Brit. (Loudon): 336 (1830). 

Kleinia haworthii DC, Prodr. 6: 338 (1838); Harvey, Fl. Cap. 3:318 (1865). 

Kleinia cana DC, Prodr. 6: 338 (1838), Harvey, Fl. Cap. 3: 319 (1865). 

K. cana was said to be allied to haworthii but with shorter leaves, ca. 17-19 
mm long (Harvey 1865). Rowley (1994) noted that the type ofK. cana DC. has 
flattened leaves which are narrowly elliptic to obovate and have rounded or pointed 
tips. Marloth in Flora of South Africa (1932: 269) remarked that difference in 
foliage "does not count" and synonymized K. cana with K. haworthii. 

Senecio quinquangulatus Sch.Bip., Flora 28: 500 (1845). 

Senecio haworthii (DC.) Sch.Bip., Flora 28: 500 (1845) (Steudel, Nomencl. Bot., 
ed. 2, 2:561, 1841,nomen). 

Because of its discoid capitula this species was treated in Kleinia by several 
authors (Haworth, De Candolle, Harvey, Marloth). Kleinia tomentosa Haw. is 
here regarded as a new name dating from 1812, not a new combination since the 
earlier potential basionym is illegitimate. 

An erect succulent sparingly branched shrublet densely silvery-white-tomentose 
throughout. Leaves 2-6-10) cm long, 6-12 mm thick, terete, fusiform to 
cylindrical, apiculate, rarely slightly flattened with a few minute lobes near the 
apex. Capitula solitary on an erect scape 8-10 cm long and with a few small 
bracts, discoid. Involucre cupshaped, phyllaries 8-9. Disc-florets 30-^10, yellow. 
Floral details as in generic description. Style branches truncate. Pappus bristles 
ca. 1 5 mm long, white. Cypsela puberulous. Chromosome no. 2n = 20 (Afzelius 
1967). -Figs. 2, 3. 

This species has a complicated nomenclatural history, which is only partly 
described by Butterfield (1954) and Rowley (1994). Very popular in cultivation 
because of its fleshy terete or fusiform leaves with a dense silvery or snow- 
white tomentum, but rarely flowering. A photograph of a flowering specimen 
in Kirstenbosch was published in Nordenstam et al. (2009) and here (Fig. 2). 
Curtis Bot. Mag. Plate 6063 was based on a flowering specimen from Sir Thomas 



66 Comp. Newsl. 50, 2012 : 



Hanbury's garden in Italy (Palazzo Orengo near Mentone), where it flowered in 

1873. 

The native habitat is in Namaqualand in the Northern Cape, e.g. in the Richtersveld, 
where the largest-leaved and whitest form has been found (cultivar 'Hans Herre' 
in Rowley 1994, 2002). Also recorded from the Great Karoo (W and E Cape), e.g. 
near Laingsburg and Camdeboo (Marloth 1932). 

3. Caputia scaposa (DC.) B. Nord. & Pelser, comb. nov. 

Basionym: Senecio scaposus DC, Prodr. 6: 403 (1838). 

Syn.: Senecio calamifolius Hook., in Curtis Bot. Mag. 69, Plate 4011 (1843). 

Stemless shrubs (var. scaposa), or erect branching short-stemmed shrub up to 0.4 
m high (var. caulescens); stems and branches pale green and somewhat downy. 
Leaves alternate, fascicled (subrosulate) in basal bunches, cylindrical, up to 12 
cm long and 0.5-1 cm thick, somewhat curved, terete and obtuse or apically 
flattened and spoon-like and lobed (var. addoensis), cobwebby-tomentose or 
white-felted, glabrescent. Capitula several or rarely solitary, terminal in a long- 
pedunculate capitulescence with reddish, sparsely tomentose, nude or shortly 
bracteate peduncles. Capitula heterogamous, radiate, yellow-flowered. Involucre 
cupshaped, calyculate; phyllaries 8-1 2, araneose. Ray-florets 7-13, female; lamina 
strap-shaped. Disc-florets hermaphrodite, corolla distally campanulate, 5-lobed. 
Floral details as in generic description. Cypselas puberulous or subglabrous. 
Chromosome no.: 2n = 20 (Afzelius 1967). - Fig. 4. 

This Eastern Cape species ranges from the Gamtoos River in Humansdorp 
eastwards to Grahamstown and Victoria East, where it grows in arid lands. It 
was introduced to Kew from a collection by Bowie (likely collected in 1820- 
1822), where it thrived anonymously for many years until it was named in 1843 
by Hooker as Senecio calamifolius. However, it was published earlier as Senecio 
scaposus in 1838 by De Candolle, who is the name-bringing author. 

Selected collections: 

CAPE PROVINCE: Uitenhage, Zwartkops River, J. F. Drege, Zeyher 2983 (BM, K, 
S), Ecklon & Zeyher 519 (K); between Bethelsdorp and Uitenhage, 27.XII.1813, 
BuRCHELL4405(herb.HooKER,K);Tambukiland,righthandsideofKeyrivierbetween 
Windvogelberg and Zwartkey, Ecklon & Zeyher 572 (S); Prope Grahamstown, 
2000 ft, IX. 1869, MacOwan 1490 (BM, K); Uitenhage, 200 ft, 13.1.1933, 
F. R. Long 895 (K); Humansdorp, 2 m. W of Gamtoos River drift on road from 
Humansdorp to Hankey, 100 ft, III. 1928, Fourcade 3623 (K); Uitenhage, on the 
Capetown road hill, 1.1840 A. Prior s.n. (K); farm Naudes Hoek near Kieskama 
River, Victoria East, H. Giffen 725 (K). 



Comp.Newsl. 50,2012 67 



Harvey recognized two varieties, viz. var. acaulis, with very short or scarcely 
any stem and var. caulescens, with a developed stem up to 40 cm and branched 
(= S. calamifolius Hook.). A third variety is var. addoensis (Rowley 1994), based 
on Senecio addoensis Compton. This differs by the flattened and lobed leaf-tips. 
Because of their morphological distinction and popularity in cultivation, the 
varieties deserve valid names as follows. 

3a. Caputia scaposa (DC.) B. Nord. & Pelser var. scaposa. 

Syn.: Senecio scaposus var. acaulis Harv., Fl. Cap. 3: 406 (1865). 

3b. Caputia scaposa (DC.) B. Nord. & Pelser var. caulescens (Harv.) B. Nord. 
& Pelser, comb. nov. 

Basionym: Senecio scaposus DC. var. caulescens Harv., Fl. Cap. 3: 406 (1865). 

Syn.: Senecio calamifolius Hook, in Curtis Bot. Mag. Plate 4011 (1843). 

3c. Caputia scaposa (DC.) B. Nord. & Pelser var. addoensis (Compton) 
B. Nord. & Pelser, comb. nov. 

Basionym: Senecio addoensis Compton, J. Bot. 72: 48 (1934). 

Syn.: Senecio scaposus DC. var. addoensis (Compton) G. D. Rowley, Cact. Succ. 
J. (Los Angeles) 62(6): 283 (1990). 

4. Caputia pyramidata (DC.) B. Nord. & Pelser, comb. nov. 

Basionym: Senecio pyramidatus DC, Prodr. 6: 402 (1838). 

A succulent short-stemmed suffrutex or shrub up to 1.8 m high with araneose- 
tomentose glabrescent stems and branches, becoming glaucous. Leaves alternate, 
mostly crowded basally, sessile, cylindrical and succulent, entire, 8-15 cm long, 
about 0.5(-l ) cm thick, acute. Capitula numerous in thyrsoid elongated or conical 
synflorescence, large, heterogamous, radiate, yellow-flowered, on bracteolate 
pedicels. Involucre hemispherical-cupshaped; phyllaries uniseriate, white- 
tomentose with brownish tips and margins. Ray-florets ca. 13 (10-14), female; 
style bilobed. Disc-florets numerous, hermaphrodite; corolla tubular, distally 
widening, 5-lobed; lobes narrowly ovate. Cypselas glabrous. Pappus bristles 
numerous, white. Chromosome no. unknown. - Fig. 5. 

Selected collections: 

CAPE PROVLNCE: Uitenhage, Bowie 3 (BM); Uitenhage, Zeyher 941 (BM); 
Zoutpanshoogde, Burke s.n. (BM); Uitenhage, XII. 1847, A. Prior s.n. (K); 
Somerset Div., mtn. above the spring at Commadagga, 6. VII. 1813, Burchell 334 1 
(K); Peddie Div., begin River Valley opposite Wooldridge, 1000 ft, 15.X.1945, 
Acocks 11852 (K); Peddie Div., near Peddie, dry Euphorbia scrub, 1800 ft, 



Comp. Newsl. 50, 2012 



7.VIII.1953, L. E. Codd 1977 (K); Algoa Bay, St. George's Strand, 20.XI.1932, 
F. R. Long 862 (K); Port Elizabeth, reed. 9.III.1906, Ethel West 20 (K); Albany, 
Pluto's Vale, 3-4 miles N of Botha's Hill, 1500 ft, X.1027, R. A. Dyer 1058 (K). 

This is a very striking species, as testified e.g. by Harvey (1865): "A very fine 
species" and Hooker in Bot. Mag. Plate 5396 stated: "perhaps among the most 
ornamental of the genus" (i.e. Senecio). 

References 

Afzelius, K. 1967. Chromosome numbers in some Senecioneae. Svensk Bot. 
Tidskr. 61: 1-9. 

Butterfield, H. M. 1954. Kleinia tomentosa and its synonyms. Cact. Slice. 
J. America 26: 153-155. 

Cullen, J., Alexander, J. C. M., Brickell, C. D., Edmondson, J. R., Green, 
P. S., Heywood, V. H., Jorgensen, P.-M., Jury, S. L., Knees, S. G., 
Maxwell, H. S., Miller, D. M., Robson, N. K. B., Walters, S. M. & P. 

F. Yeo 2000. The European Garden Flora. Vol. 6, Dicotyledons (Part IV). 
Cambridge Univ. Press, Cambridge, U. K. 

De Candolle, A. P. 1838. Prodromus systematis naturalis regni vegetabilis. Vol. 
6. Treuttel et Wiirtz, Paris. 

Eggli, U. (ed.) 2002. Illustrated handbook of succulent plants: Dicotyledons. 545 
+ lxiv pp. Springer- Verlag, Berlin, Heidelberg, New York. 

Harvey, W. H. 1 865. Compositae. Pp. 44-530 in: Harvey, W. H. & O. W. Sonder, 
Flora capensis 3. Hodges, Smith & Co., Dublin; I. C. Juta, Capetown. 
Reprint ed., 1894; Lovell Reeve & Co., Covent Garden, London. 

Hilliard, O. M. 1977. Compositae in Natal. University of Natal Press, 
Pietermaritzburg. 

Holmgren, P. K., Holmgren, N. H. & L. C. Barnett (eds). 1990. Index 
Herbariorum. Part 1: The Herbaria of the World. Ed. 8. Regnum Vegetabile 
vol. 120. New York Botanical Garden, Bronx, New York. 

Hutchinson, J. 1923. Senecio Medley- Woodii. Plate 83 in: Pole Evans, I. B. (ed.), 
The Flowering Plants of Africa. Vol. III. L. Reeve, London; Speciality Press 
S. A-, Johannesburg, Capetown. 

Marloth, R. 1932. The flora of South Africa, Vol. 3, 2. Darter Bros. & Co., 
Capetown; Wheldon & Wesley, London. 



Comp. Newsl. 50, 2012 69 



Nordenstam, B. 2007. Tribe Senecioneae Cass. Pp. 208-241 in: Kubitzki, K. 
(ed.), The families and genera of vascular plants Vol. 8, Flowering plants: 
Eudicots, Asterales (Kadereit, J. W. & C. Jeffrey, eds.). Springer, Berlin. 

Nordenstam, B., Pelser, P. B., Kadereit, J. W. & L. E. Watson 2009. Chapter 
34. Senecioneae. Pp. 503-525 in: Funk, V. A., Susanna, A., Stuessy, T. 
F. & R. J. Bayer (eds.), Systematics, Evolution, and Biogeography of 
Compositae. IAPT, Vienna. 

Pelser, P. B., Nordenstam, B., Kadereit, J. W. & L. E. Watson 2007. An 
ITS phylogeny of tribe Senecioneae (Asteraceae) and a new delimitation of 
Senecio L. Taxon 56: 1077-1104. 

Pelser, P. B., Kennedy, A. H., Tepe, E. J., Shidler, J. B„ Nordenstam, B., 
Kadereit, J. W. & L. E. Watson 2010. Patterns and causes of incongruence 
between plastid and nuclear Senecioneae (Asteraceae) phylogenies. Amer. 
J. Bot. 97(5): 856-873. 

Rowley, G. D. 1994. Succulent Compositae. Strawberry Press, Mill Valley, 
California. 

Rowley, G. D. 2002. Senecio. Pp. 29^13 in: Eggli, U. (ed.) 2002. Illustrated 
handbook of succulent plants: Dicotyledons. 545 + lxiv pp. Springer- Verlag, 
Berlin, Heidelberg, New York. 



70 Comp. Newsl. 50, 2012 

Crassothonna B. Nord., a new African genus of 
succulent Compositae-Senecioneae 

Bertil Nordenstam 



Department of Phanerogamic Botany, Swedish Museum of Natural History 

P. O. Box 50007, SE-10405 Stockholm, Sweden 

bertil.nordenstam@nrm.se 



Abstract 

The new genus Crassothonna B. Nord. is described with 13 species transferred 
from the large and polymorphic genus Othonna L. Crassothonna species are 
perennial herbs, shrublets or shrubs with terete succulent leaves. The genus is 
mainly South African with a centre in Little Namaqualand and adjacent parts of 
the Western Cape, but is also represented by a few species in the eastern Cape 
Province, Natal and southern Namibia. Several species belong to the Gariep 
floristic element. 

Introduction 

The genus Othonna L. (Compositae-Senecioneae) comprises about 120 species 
(Nordenstam 2007) with a marked centre in southern Africa. The taxonomy 
of this genus is notoriously difficult, and further revisional work remains to be 
done especially on the specific level. On the generic level one major change is 
necessary, viz. to remove a group of highly succulent terete-leaved taxa out of 
the genus. They are here recognized as a new genus Crassothonna B. Nord. with 
(provisionally) 13 species. The remaining Othonna s.str. comprises tuberous 
herbs and shrubby perennials with quite a variety of life- forms and adaptations to 
extreme climatic conditions. 

Molecular data support the segregation of this group from Othonna, although only 
a few species have been investigated so far. However, it has been demonstrated that 
O. sedifolia and carnosa (now C. cacalioides) form a clade closer to Gymnodiscus 
Less, (a small genus of annual herbs) than to the core of Othonna (Devos et al. 
2010, Nordenstam et al. 2009, Pelser et al. 2010). 



Comp.Newsl. 50,2012 71 



Crassothonna B. Nord., gen. nov. 

Glabrous shrubs or shrublets or perennial herbs. Stems terete, thin to thickened, 
sometimes bottle-shaped and carnose, little- to much-branched. Leaves alternate, 
simple, terete, subterete or ovoid to fusiform, highly succulent, green or glaucous 
or pruinose or minutely papillose, sessile or shortly petiolate, apically rounded 
to obtuse, sometimes mucronate. Capitula heterogamous, radiate or disciform, 
terminal, pedunculate on thin and sometimes long peduncles with one or a few 
minute bracts, solitary or few to several or many in corymbose synflorescence. 
Involucre soft, campanulate (not cup-shaped), phyllaries uniseriate, 5-8, basally 
shortly connate, herbaceous, 3-5-veined, with membranaceous margins.. 
Receptacle flat or somewhat convex, glabrous, minutely alveolate. Ray-florets 
female, fertile, tube shortly cylindrical, lamina strap-shaped, yellow or more 
rarely white, sometimes tubular and eligulate. Style bifurcate, truncate or obtuse, 
with short apical sweeping-hairs. Cypselas fusiform-oblong-obovoid, 2-5 mm 
long, 1-3 mm wide, veined or ribbed, glabrous or puberulous or covered with a 
dense silvery-white tomentum of appressed trichomes becoming mucilaginous 
when soaked. Pappus bristles numerous, minutely barbellate, white, persistent or 
caducous. Disc-florets hermaphrodite but often female-sterile (functionally male), 
with style in central florets undivided with a conical tip, but in some taxa marginal 
disc-floret styles fertile, bifurcate with separate stigmatic areas on the inside of 
style branches and tip shortly conical or subtruncate with short sweeping-hairs; 
corolla tubular-campanulate, 5-lobed, lobes midveined, sometimes only faintly 
so. Anthers basally obtuse, ecaudate, endothecium radial; apical appendage flat, 
ovate-oblong; filament collar balustriform. Cypselas, usually not developing, 
narrowly oblong, glabrous or puberulous. Pappus bristles few to several, white, 
caducous. 

Type: Crassothonna cylindrica (Lam.) B. Nord. 

13 species recognized here, South Africa, Namibia. A few more taxa remain to 
be described, and a revision of the genus is in preparation. The members of the 
genus have a characteristic look. The terete succulent leaves are usually long and 
uniformly thick, but variable in size and shape. They may be short and almost 
grape-like as in C. clavifolia or small and somewhat larva-like as in C. sedifolia, 
and in C. opima very large and sausage-like. 

Most species are diploid with 2n=20. This number has been counted in C. alba, 
cacalioides (2 collections), capensis (2 collections), clavifolia, cylindrica, 
floribunda, opima, patula, protecta (6 collections), sedifolia (3 collections) and 
sparsiflora (Afzelius 1924, 1967, Czeika 1957, Nordenstam 1967, 1969a, 1971, 
Ornduff et al. 1967, Ratler & Milne 1973). Only C. rechingeri differs by the 
hexaploid number 2n=60 (Nordenstam 1971 ). 



72 Comp. Newsl. 50, 2012 

1. Crassothonna alba (Compton) B. Nord., comb. nov. 

Basionym: Othonna alba Compton, Trans. Roy. Soc. South Africa 19: 321 (1931). 

Similar to C. cylindrica and C. cacalioides but easily recognized by the white 
rays. Rowley ( 1 994), also in Eggli (2002), did not recognize it as a distinct species 
but cited it in synonymy of Othonna carnosa Less., which is now Crassothonna 
cacalioides. The status of this white-rayed taxon will be further investigated. It 
has been recorded from the Western Cape, in karroo, but is nowhere common or 
abundant. 

2. Crassothonna cacalioides (L.f.) B. Nord., comb. nov. 

Basionym: Cineraria cacalioides L.f, Suppl. PL: 374 (1782). 

Syn.: Othonna carnosa Less., Syn. Gen. Compos.: 88 (1832). 

Lessing published the new name Othonna carnosa for a taxon cited as Cineraria 
cacalioides Thunb. and for the obvious reason that the specific epithet was not 
available under Othonna (O. cacalioides L. fil. is a quite different species and a true 
Othonna; cf Nordenstam 1969b). This nomenclature was adopted by Harvey in 
Flora capensis (Harvey 1865). However, under Crassothonna the earliest epithet 
cacalioides is available, so this species has to change its name completely - from 
Othonna carnosa to Crassothonna cacalioides. The white-rayed related taxon 
from the Western Cape is distinguished as C. alba (cf. above), and a rayless taxon 
in KwaZulu-Natal Province is recognized as C. discoidea (cf. below). 

Coastal areas from Humansdorp in the Eastern Cape north to KwaZulu-Natal 
border. 

3. Crassothonna capensis (L. H. Bailey) B. Nord., comb. nov. 

Basionym: Othonna capensis L. H. Bailey, Cycl. Amer. Hort.: 1180 (1901). 

Syn.: Othonna crassifolia Harv., Fl. Cap. 3: 336 (1865), non O. crassifolia L., 
Syst. Nat. ed. 12, 2: 579; Mant. PL: 1 18 (1767) = Othonna othonnites (L.) Druce. 

Very popular and common in cultivation, and unfortunately sometimes still under 
the invalid name from Harvey's Flora capensis, viz. Othonna crassifolia Harv. 

The plants in cultivation easily form green mats of succulent leaves and a profusion 
of yellow flowerheads, which makes it a favourite in rock gardens or green-houses 
in many parts of the world. With reference to the leaf-shape it is often called 
'Little Pickles'. 

Distributed in the Western Cape. 



Comp.Newsl. 50,2012 73 



4. Crassothonna clavifolia (Marl.) B. Nord., comb. nov. 

Basionym: Othonna clavifolia Marloth, Trans. Roy. Soc. South Africa 2: 38 
(1910). 

Short thickened stems and leaves grape-like or ovoid-fusiform and mucronate. 

This species is endemic to the area around the lower Orange River, i.e. 
southernmost Namibia and the Richtersveld in northern Namaqualand, a region 
phytogeographically known as the Gariep Centre, home of the Gariep floristic 
element (Nordenstam 1966, 1969c, van Wyk & Smith 2001). 

The species is sought after by succulent growers and makes a nice impression with 
its low habit and grape- or olive-like leaves in small bunches. 

5. Crasssothonna cylindrica (Lam.) B. Nord., comb. nov. 

Basionym: Cacalia cylindrica Lam., Encycl. Meth. Bot.l : 529 (1785). 

Syn.: Othonna cylindrica (Lam.) DC, Prodr. 6: 477 (1838). 

A branching erect shrub up to one m high, with elongate terete leaves and several 
(mostly 2-4) yellow-flowered capitula in small synflorescences. 

This is a shrubby species widely distributed in the Western Cape north to southern 
Namibia. The succulence of the leaves has been questioned (cf. Rowley 1994: 
170), but there is no doubt that the leaves are terete and succulent, like in all 
congeners. 

6. Crassothonna discoidea (Oliv. in Hook.) B.Nord., comb, et stat.nov. 

Basionym: Othonna carnosa var. discoidea Oliv. in Hook. Icon. Plant. 18 t. 1713 
(1887-1888). 

This is sufficiently distinct from C. cacalioides to be regarded as a separate species 
(cf. Hilliard 1 977, who expressed the same opinion). Apart from the disciform (not 
discoid, in spite of the epithet) capitula it has a distinctive habit with rather large 
and long leaves (up to 15 cm long and 1 cm wide), many-flowered capitulescences 
with long peduncles, and other characteristics to be discussed in my forthcoming 
revision of the genus. It seems to be confined to KwaZulu-Natal, where it is found 
in coastal areas, forming colonies in sand dunes or sandy grassland. 

7. Crassothonna floribunda (Schltr) B. Nord., comb. nov. 

Basionym: Othonna floribunda Schltr, Bot. Jahrb. Syst. 27(1-2): 214 (1899). 

This is close to C. cylindrica with which it is partly sympatric, distributed in 
the Western Cape, from Malmesbury northwards into Namaqualand, but has not 
yet been recorded from Namibia. It is more richly flowering (= 'floribund') than 



74 Comp. Newsl. 50, 2012 

C. cylindrica with rays deeper yellow or even orange-coloured. 

8. Crassothonna opima (Merxm.) B. Nord., comb. nov. 

Basionym: Othonna opima Merxm., Mitt. Bot. Staatssamml. Munchen 5: 636 
(1965). 

A robust erect branching shrub up to one m high with thickish stems and branches. 
Leaves very long and thick, terete, sausage-like, up to 15 cm long and 1.5 cm 
thick. 

Perhaps the most striking species in the genus, because of its large sausage- 
like leaves. It is confined to the Richtersveld in northern Namaqualand and 
southernmost Namibia, i.e. a typical Gariep element. It is hybridizing occasionally 
with one or two other species of Crassothonna, viz. C. cylindrica and C.floribunda 
(cf. also Rowley 1994: 185). I have seen and collected intermediate specimens 
growing with the presumed parent species in parts of the Richtersveld. 

9. Crassothonna patula (Schltr) B. Nord., comb. nov. 

Basionym: Othonna patula Schltr, J. Bot. 36: 26 (1< 



A stemless herbaceous succulent with stolons, forming new plants from rooting 
nodes. 

This was described from the Eastern Cape, but is apparently distributed more 
widely with scattered occurrencies also in the Western Cape and Namaqualand. 
The stoloniferous habit is shared with C. rechingeri (cf. below). The latter is a 
hexaploid rayless species, whereas C. patula is a radiate diploid with 2n=20, 
counted on plants originating from Vanrhynsdorp Div., 1 5 miles E of Vanrhynsdorp 

(NORDENSTAM 1971). 

10. Crassothonna protecta (Dinter) B. Nord., comb. nov. 

Basionym: Othonna protecta Dinter, Repert. Spec. Nov. Regni Veg. 19: 141 
(1923). 

Syn.: Othonna crassicaulis Compton, Trans. Roy. Soc. South Africa 19: 322 
(1931). 

A characteristic species with a bottle-shaped or sausage-like swollen short stem 
up to 15-20 cm high, little-branched with weak branches. Leaves subterete and 
succulent, but often somewhat flattened or grooved on the upper side, rather 
narrow and elongate, apically mucronate; leaf-axils woolly. Ray-florets numerous 
(13—22), with light yellow or greenish lamina, often becoming rolled back. 

The species is distributed in arid areas of the Western and Northern Cape from the 
Witteberg and Karoo Poort through Namaqualand into southern Namibia. 



Comp.Newsl. 50,2012 75 



11. Crassothonna rechingeri (B. Nord.) B. Nord., comb. nov. 

Basionym: Othonna rechingeri B. Nord., Ann. Naturhist. Mus. Wien 75: 139 
(1971 publ. 1972). 

This one of the few rayless species of the genus. The stoloniferous habit is shared 
with C. patula, which is a diploid with radiate capitula. C. rechingeri is hexaploid 
with 2n=60 (Nordenstam 1971). The species is easily cultivated and tends to 
spread to other pots by the rooting tuberous nodes formed on the runners. 

The species may be confined to the Northern Cape (type locality in Calvinia Div.). 

12. Crassothonna sedifolia (DC.) B. Nord., comb. nov. 

Basionym: Othonna sedifolia DC, Prodr. 6: 479 (1838) [1837 publ. early Jan 
1838]. 

A small erect branching shrub up to 50-60 cm high with mostly single capitula. 
Leaves short and thick, obovoid or terete, less than 1 cm long and shortly petiolate 
to subsessile, smooth or minutely papillose. Rays pale or richly yellow. 

There is some interesting variation within this species especially in leaf shape 
and texture, and it is possible that Othonna papillosa Dtr may be recognized as a 
separate taxon. Further studies on this issue are ongoing. 

Fairly widespread in the Western Cape and Namaqualand into southern Namibia. 

13. Crassothonna sparsiflora (S. Moore) B. Nord., comb. nov. 

Basionym: Euryops sparsiflorus S. Moore, Bull. Herb. Boissier Ser. 2, 4: 1023 
(1904). 

Othonna sparsiflora (S. Moore ) B. Nord., Mitt. Bot. Staatssamml. Miinchen 4: 
125(1961). 

This species was first described as a member of Euryops, but moved to Othonna 
by Nordenstam (1961). It is similar to C. cylindrica and C. cacalioides but easily 
recognized on its disciform heads with tubular-campanulate hermaphroditic 
marginal florets. The plant is a branching shrub ca. 0.5 to 0.8 m tall and the 
capitula are borne singly on peduncles up to 7 cm long. 

Restricted to the Richtersveld in northern Namaqualand and southern Namibia, 
i.e. a typical Gariep element (cf. spp. 4 & 8). 



76 Comp. Newsl. 50, 2012 



References 

Afzelius, K. 1924. Embryologische und zytologische Studien in Senecio und 
verwandten Gattungen. Acta Hort. Berg. 8: 123-219. 

Afzelius, K. 1967. Chromosome numbers in some Senecioneae. Svensk Bot. 
Tidskr. 61: 1-9. 

Czeika, G. 1957. Strukturveranderungen endopolyploider Ruhekerne in 
Zusammenhang mit Wechseln der Bundelung der Tochter-chromosomen 
und karyologisch-anatomische Untersuchungen an Sukkulenten. Osterr. 
Bot. Zeitschr. 103: 536-566. 

Devos, N., Nordenstam, B., Mucfna, L. & N. P. Barker 2010. A multi-locus 
phylogeny of Euryops (Asteraceae, Senecioneae) augments support for the 
"Cape to Cairo" hypothesis of floral migrations in Africa. Taxon 59: 57—67. 

Eggli, U. (ed.) 2002. Illustrated handbook of succulent plants: Dicotyledons. 545 
+ Ixiv pp. Springer- Verlag, Berlin, Heidelberg, New York 

Harvey, W. H. 1 865. Compositae. Pp. 44-530 in: Harvey, W. H. & O. W. Sonder, 
Flora capensis 3. Hodges, Smith & Co.. Dublin; I. C. Juta, Capetown. 
Reprint ed., 1894; Lovell Reeve & Co., London. 

Hilliard, O. M. 1977. Compositae in Natal. University of Natal Press, 
Pietermaritzburg. 

Nordenstam, B. 1961. Die Gattungszugehorigkeit von Euryops sparsiflorus 
S. Moore. Mitt. Bot. Staatssamml. Munchen 4: 125-126. 

Nordenstam, B. 1966. Ewyops in South West Africa. Bot. Notiser 119: 475-485. 

Nordenstam, B. 1967. Chromosome numbers in Othonna (Compositae). Bot. 
Notiser 120: 297-304. 

Nordenstam, B. 1969a. Chromosome studies on South African vascular plants. 
Bot. Notiser 122:398^08. 

Nordenstam, B. 1969b. Othonna cacalioides. Flow. Plants Afr., Plate 1572. 

Nordenstam, B. 1969c. Phytogeography of the genus Euryops (Compositae). 
Opera Bot. 23, 77 pp. 

Nordenstam, B. 1971. Othonna rechingeri B. Nord., spec, nova, a hexaploid 
succulent from South Africa. Ann. Naturhistor. Mus. Wien 75: 139-142. 

Nordenstam, B. 2007. Tribe Senecioneae Cass. Pp. 208-241 in: Kubitzki, K. 
(ed.), The families and genera of vascular plants Vol. 8, Flowering plants: 
Eudicots, Asterales (Kadereit, J. W. & C. Jeffrey, eds.). Springer, Berlin. 



Comp.Newsl. 50,2012 77 



Nordenstam, B., Clark, V. R., Devos, N. & N. P. Barker 2009. Two new 
species of Ewyops (Asteraceae: Senecioneae) from the Sneeuberg, Eastern 
Cape Province, South Africa. S. Afr J. Bot. 75: 145-152. 

Ornduff, R., Mosquin, Th., Kyhos, D. W. & P. H. Raven 1967. Chromosome 
numbers in Compositae. VI. Senecioneae. II. Amen J. Bot. 54: 205-213. 

Pelser, P. B., Kennedy, A. H., Tepe, E. J., Shidler, J. B., Nordenstam, B., 
Kadereit, J. W. & L. E. Watson 2010. Patterns and causes of incongruence 
between plastid and nuclear Senecioneae (Asteraceae) phylogenies. Amer. 
J. Bot. 97(5): 856-873. 

Ratler, J. A. & C. Milne 1973. Some Angioperm chromosome numbers. Notes 
Roy. Bot. Gard. Edinb. 32: 429^138. 

Rowley, G. D. 1994. Succulent Compositae. Strawberry Press, Mill Valley, 
California. 

van Wyk, A. E. & G. Smith 2001. Regions ofFloristic Endemism in Southern 
Africa. Umdaus, Hatfield, S.A. 



78 Comp. Newsl. 50, 2012 

Diversity of trichomes from mature cypselar 
surface of some taxa from the basal tribes of 

Compositae 

Sobhan Kr. Mukherjee i & Bertil Nordenstam 2 



'Taxonomy and Biosystematics Laboratory 
Department of Botany, University of Kalyani 

Kalyani, Nadia-741 235. W. B., India 
sobhankr@gmail.com; sobhankr@yahoo.com 

department of Phanerogamic Botany 

Swedish Museum of Natural History 

SE-104 05 Stockholm, Sweden 

bertil.nordenstam@nrm.se 



Abstract 

Structure and distribution of trichomes are important from the taxonomic point 
of view. The distribution and structure of trichomes appear to be genetically 
controlled and are more or less stable and have paramount taxonomic significance. 

It has been observed by many workers that trichomes from the vegetative as well 
as reproductive parts can be used successfully for the delimitation of genera and 
species within the family Compositae ( Asteraceae). However, broad comparative 
studies of cypselar trichomes are scarce or lacking. The present study indicates 
that trichome structure is highly variable. Trichomes from the outer surface of 
mature cypselas of 44 genera and 7 1 species belonging to 7 tribes of the four 
subfamilies Mutisioideae. Carduoideae, Pertyoideae and Cichorioideae were 
studied by light microscopy and SEM. Number of studied genera and species in 
each tribe is indicated in parentheses: Mutisieae (1/3), Dicomeae (1/1), Cardueae 
(9/17), Pertyeae (1/3), Cichorieae (14/23), Arctoteae (3/3) and Vernonieae (12/21). 
Both glandular and non glandular trichomes are divided into seven major types. 

Presence of 'twin' or 'duplex' cypsela hairs is one of the characteristic features 
of the family Compositae and is prevalent in the tribes Cichorieae, Dicomeae, 
Pertyeae, Mutisieae and Vernonieae. This type of trichomes is less common in 
the tribes Arctoteae and Cardueae. Distribution and structure of trichomes have 
taxonomic significance at the generic and infrageneric level. However, their value 



Comp. Newsl. 50, 2012 79 



as taxonomic markers will be greatly increased when combined with other cypsela 
characters such as structure of carpopodium, stylopodium and pappus, as well as 
surface features of cypselar wall, etc. Certain taxa have unique types of trichomes 
regarded as apomorphies. Fusiform twin hairs have been noted in Gerbera 
piloselloides, whereas papillose twin hairs were noticed in Actites megalocarpa. 
Sometimes papillate hairs united to form a scale-like structure e.g. in Crepis and 
Hypochaeris. Multicellular non-glandular acroscopic hairs have been noticed in 
Echinops sphaerocephalus . In Carlina, glandular twin hairs were observed. 

Keywords: Trichomes, twin hairs, mature cypselas, Compositae. 

Introduction 

The Compositae (Asteraceae) family is nested in the order Asterales of Campanulids 
according to the APG III classification (The Angiosperm Phylogeny Group 2009). 
The family has largest number of accepted species, ca. 24,000, among angiosperm 
plant families and is distributed throughout the globe except Antarctica (Funk 
et al. 2009, Funk 2010). The genera number 1600-1700. All available evidence 
indicates that the family is monophyletic (e. g.. Small 1919, Bremer 1987, Jansen 
& Palmer 1987, 1996, Bremer et al. 1992, Hansen 1990, 1991a, b). 

After the long outdated classifications of Bentham (1873 a, b) and Hoffmann 
(1890), the new classification of the Compositae by Bremer (1994) was the first 
approach at a modern phylogenetic analysis of this family. Bremer's classification 
was morphologically based, but with subsequently rapidly developing molecular 
data a new picture of the family phylogeny emerged, first summarized by Kadereit 
& Jeffrey (2007) and their associates. This modern approach culminated in the 
seminal and epoch-making work of Funk et al. (2009). At present, the basal groups 
of Compositae are recognized as subfamilies like Barnadesioideae, Mutisioideae, 
Hyalideae, Wunderlichieae, Carduoideae, Pertyoideae and Cichorioideae. The 
present study includes some members of four basal subfamilies (Mutisioideae, 
Carduoideae, Pertyoideae and Cichorioideae). 

Trichomes or hairs have long been of much importance in systematic investigation 
of angiosperms. Anatomically, a trichome or hair is an epidermal outgrowth of 
diverse form, structure and function (Uphof 1962, Esau 1965, Fahn 1986, 1988). 
Despite the great variety of systems proposed for the classification of trichome 
types, they are basically classified as either glandular with a secretary function 
or as covering hairs (non-glandular) without a secretary function (Cutter 1978). 
Hairs originate from epidermal cells and develop on the outer surface of various 
plant organs (Werker 2000). The hairs offer a rich field of investigation for a 
micro-morphologist because of their common occurrence and great diversity, 



80 Comp. Newsl. 50, 2012 



simplicity in structure and easy availability for observation due to their superficial 
position (Sahu 1982 a, b, c, 1983). 

The types of hairs (epidermal outgrowths) have been reviewed and classified in 
early works, such as by De Bary (1884) who recognized four major categories, 
viz., (i) Bladders, (ii) Hairs, (iii) Scales and (iv) Shaggy hairs. De Bary (I.e.) also 
noted that the presence of hairs in plant organs is a universal character in vascular 
plants. Later Cowan (1950) merged the 'Shaggy hairs' into the category of 'hairs' 
and various other categorizations of trichome type have been put forward. Similar 
approaches have been made in various plant families. For example, Goebel ( 1 900) 
was the first to discuss the significance and relationships of trichomes in the tribe 
Rhinantheae of the family Scrophulariaceae. 

One pioneer in the study of role and importance of hairs in taxonomy and their 
phylogenetic evolution was Solerader (1908) in his work entitled "Systematic 
Anatomy of the Dicotyledons". He again divided the hairs into two major 
categories, viz. (i) the clothing hairs and (ii) the glandular hairs. Solerader 
(op.c.) described the hairs present on the ovary of Spilanthes oleracea (now 
Acmella oleracea) as follows: "they consist of two long hair cells which are 
jointed together lengthwise along one side, but diverse apically so as to resemble 
swallows tail." Such types of hair are characteristic of the ovary or cypsela 
(achene) in the family Compositae (Ramayya 1962 a, b, c) and these hairs were 
designated variously; e.g., 'Zwillingshaare' (Kraus 1866, Hess 1938), translated 
to 'twin hairs' (e.g., Vaughan 1970, Grau 1977, Dittrich 1977), or 'biseriate 
forked hair' (Narayana 1979), simply 'achenial hair' (Small 1919, Hess 1938, 
Ramayya 1962 a, b, c, 1963, 1969, 1972, Manilal 1963, Sahu 1976, 1978, 1979, 
1980, 1982 a, b, 1983, 1984, Freire etal. 2002), or sometimes 'duplex hair' (e.g., 
Macloskie 1883, Nichols 1893,Nordenstam 1977). Primarily a twin hair consists 
of two triangular or rectangular and usually short basal cells, completely or partly 
united with each other along their longitudinal walls (Sancho & Kaunas 2002). 
Nordenstam (1968 b) described the twin hairs of Ewyops as consisting of two 
parallel, short to elongate cells plus a smaller basal cell, i.e. three cells altogether. 

Early records of the hairs of Compositae are usually brief and sometimes 
fragmentary, e.g. as reported by Nichols (1893), Lloyd (1901), Kupfer (1903), 
Cavillier(1907, 1911), Holm (1908, 1913, 1917 & 1926), Sabnis (1921), Briquet 
(1916, 1930), Diettert (1938), Artschwager (1943), Volle & Hetzger (1949) 
and Mirashi (1955, 1956). Later more detailed accounts have been made such as 
the descriptions of glandular hairs of the subtribe Madinae presented by Carlquist 
(1958, 1959 a, b, c). 

Another significant contribution regarding the hairs of Compositae has been 
documented by Ramayya (1962 a, b, c, 1963 &1969). He described the structure, 



Comp. Newsl. 50, 2012 81 



variation and distribution of 35 types of hairs in some Compositae and their 
mode of development. He also recognized four distinct "Systems of hairs" in this 
family, viz., (i) Filiform Trichome System; (ii) Macroform Trichome System; (iii) 
M-Multiseriate Trichome System and (iv) P-Multiseriate Trichome System. 

Metcalfe & Chalk (1950, 1979, 1989) have stated that trichomes possess 
taxonomic value, that both glandular and non-glandular hairs are present in 
the Compositae, and that the non-glandular hairs are of various morphological 
types, whereas the glandular hairs are more or less homogeneous with a uniform 
structure. 

Structures of hairs are greatly variable among different taxa. For example, Spring 
(2000) has described more than 300 forms of plant trichomes. 

Many workers have contributed on the systematic value of hairs in the genus 
Vernonia s.lat, like Hunter & Austin (1967) and Faust & Jones (1973) in North 
American species, and Narayana (1979) on 15 South Indian species. In the 
latter 18 types of hairs were noted by Narayana (I.e.), out of which three types 
(biseriate vesicular glandular hair, biseriate non-forked hair and biseriate forked) 
were found on the ovary wall. Twenty Indian species of Vernonia were studied 
by Sahu (1984), who mentioned that the frequency distribution of cypselar (= 
achenial) hairs is highest (57%) among all the hairs observed. Five among the six 
cypselar hair types noted are of glandular type, viz. (i) biseriate glandular hair, 
(ii) biseriate vesicular glandular hair, (iii) biseriate capitate glandular hair, (iv) 
biseriate vesicular capitate glandular hair, and (v) multiseriate glandular hair, and 
one (vi) non-glandular hair type. 

Nordenstam (1968b, 1977, 1978, 2006a, 2007) noted that the duplex hairs are 
widespread in the tribe Senecioneae but not in Calenduleae. He found that such 
hairs show various modifications and occasionally (e.g. in species of Euryops, 
Mesogramma, Bolandia, Dauresia, Lomanthus) exude mucilage when treated in 
water (Nordenstam & Pelser 2005, 2009). Similar type of duplex hairs has been 
observed by Macloskie (1883), James (1883), Nichols (1893), Drury & Watson 
(1965) and Sahu (1983), mainly in the Senecioneae, e.g. species of Senecio, 
Othonna, and Euryops. Macloskie (1883) and Sahu (1976, 1978, 1979, 1980, 
1982 a, b) also observed similar trichomes in other taxa of Compositae. 

King & Robinson (e.g., 1969, 1970, 1979) have used the hair structure along 
with other epidermal features in determining many generic circumscriptions in 
the Compositae, especially in the Eupatorieae. 

In the systematic review of Cynareae, Dittrich (1977) reported that cypselas 
in subtribe Echinopsinae (Echinops and Acantholepis) are densely covered by 
multicellular stiff hairs. In all members of the subtribe Carlininae, cypsela hairs 



82 Comp. Newsl. 50, 2012 



are 'twin hair' type. Members of the Cardueae have been studied in some detail 
by Petit (1987, 1997). 

Napp-Zinn & Eble (1980) studied the glandular and non glandular hairs of 
20 genera of the Anthemideae. On the basis of differences of glandular hairs, 
three species of Artemisia are clearly separated, i.e., A. nova, A. arbuscula and 
A. tridentata (Kelsey & Shafizadeh 1980, Kelsey 1984). 

The types of hairs may be considered as valuable accessory characters in plant 
taxonomy, particularly at infrageneric levels (Stebbins 1953, Faust & Jones 1973, 
Napp-Zinn & Eble 1980, Sahu 1978, 1979, 1980, 1982 a, b, 1984, Hoot 1991, 
Korolyuk 1997). 

Sancho & Kaunas (2002) have shown the presence of typical twin type of hairs 
from the corollas of Mutisieae. In addition to that corollas also bear four other 
types of hairs. They have also indicated that the ontogenies of cypselar twin hairs 
and twin hairs of corollas were identical. 

Nordenstam (1968a, 1978, 2006 b) and Lundin (2006) have pointed out that some 
members of the tribe Senecioneae possess more complex, stellate or substellate 
types of hairs, e.g., Aequatorium B. Nord. and Nordenstamia Lundin and rarely 
in Euryops, although not on the cypselas, which are glabrous (or glandular- 
puberulous). Substellate or T-shaped hairs occur also in Dress lerothamnus 
H. Rob. of the same tribe (Robinson 1989). 

Microcharacters of glandular hairs from 34 species belonging to 7 tribes of 
the subfamily Asteroideae, have been studied by Ciccarelli et al. (2007) who 
discussed the usefulness of the glandular hairs of the ovary. 

Leaves of the industrial oilseed Vernonia galamensis ssp. galamensis var. 
ethiopica Gilbert, contain glandular hairs, which are actually 10-celled peltate 
biseriate glandular hairs (cf. also below). Along with these hairs this taxon has 
awl-shaped glandular hairs and non-glandular hairs (Favi et al. 2008). 

The detailed structure of trichomes from stem and leaves of 135 species belonging 
to 53 genera in tribe Lactuceae was studied by Krak & Mraz (2008). They 
recognized eight types and several subtypes, and they have focused on the utility 
of hairs in the infratribal classification with the help of both light and scanning 
electron microscopy. 

Andreucci et al. (2008) studied critically the histochemistry and morphology of 
multicellular biseriate glandular hairs of Matricaria chamomilla. These hairs are 
composed of 10 cells, viz. two basal cells, two peduncle cells and a secretary head 
containing six cells. Ten-celled biseriate glandular hairs also have been reported by 
Monteiro et al. (2001) from the leaves of Stevia rebaudiana (tribe Eupatorieae). 



Comp.Newsl. 50, 2012 83 



Similar type of stalked glandular hair and two types of non-glandular hairs have 
been noted from the cypsela of Stevia rebaudiana by Cornara et al. (2001). 

Adedeji (2004) and Adedeji & Jewoola (2008) stated that hairs can be fruitfully 
used for the delimitation of genera within the family Compositae and they have 
mentioned different types of glandular and non-glandular hairs. The genus 
Vernonia can be delimited from other genera by the presence of T-shaped hairs 
and Chromolaena has amoeboid-shaped hairs. 

Diversity of leaf hairs and phylogenetic relationships on the basis of such hairs 
in different taxa of Artemisia have been discussed by Hayat et al. (2009a, b, c ) 
using both light microscopy and scanning electron microscopy. Eight major types 
of hairs were distinguished in their study. 

Robinson (2009) has correlated some hair characters with their chemical 
components. Glandular hairs may have different types of defensive components, 
viz. sesquiterpene lactones, monoterpenoids, gelatinous material with clerodane 
and labdane derivatives, which are usually common in Eupatorieae and Heliantheae. 
These two tribes are also characterized by the absence of T-shaped hairs. He 
has also emphasized the use of non-glandular hairs in different degrees. Hair 
characters support the separation of Critoniopsis Sch.Bip. from Tephrothamnus 
Sch.Bip. and Eremosis (DC.) Gleason as proposed by Keeley et al. (2007) on the 
basis of DNA studies. 

Adding to the great variation in cypselar hairs, the tribe Gochnatieae (previously 
in Mutisieae) has three types of hairs, viz., two-armed, obliquely septate, and 
biseriate glandular (Freire et al. 2002). Ventosa & Herrera (2011 a, b) report 
the presence of biseriate glandular hairs with vesicle and simple biseriate non- 
glandular hairs on the cypselas of several species of Anastraphia D. Don (formerly 
in genus Gochnatia). 

The present study deals with the cypselas of 44 genera and 71 species of 7 
tribes (Mutisieae, Dicomeae, Cardueae, Pertyeae, Cichorieae, Arctoteae and 
Vernonieae) under four subfamilies (Mutisioideae, Carduoideae, Pertyoideae 
and Cichorioideae). We thus briefly survey and summarize the variation of hairs 
from the mature cypselas belonging to the subfamily Cichorioideae sensu Bremer 
(1994), or the basal subfamilies and tribes sensu Panero & Funk (2002), Funk 
etal. (2009) and Funk (2010). 

The objectives of this study are as follows: 

i) to elucidate the presence or absence of hairs on cypselar surface. 
ii) to present the types of hairs and location of hairs on the cypselar surface. 
iii) to describe the comparative morphological features of cypselar hairs in 
different genera and species. 



84 Comp. Newsl. 50, 2012 



iv) to facilitate an accurate and rapid identification of taxa along with other 

cypselar features. 
v) to verify the potential usefulness of cypselar hairs for taxonomic or 

phylogenetic studies. 
vi) to investigate affinities among the analyzed taxa based on the distribution of 

hairs types. 

Since micro-morphological features of hairs from different plant parts play such 
a significant role in plant taxonomy, particularly at the generic and specific levels, 
many plant anatomists and taxonomists have been attracted and fascinated by this 
subject and tried to prove the relationships of taxa on the basis of hair characters. 
Only a fraction of the rich literature has been summarized here. But knowledge 
about cypselar hairs and their taxonomic significance is still incomplete. Therefore, 
some additional information based on both light microscopy (LM) and scanning 
electron microscopy (SEM) is provided here. 

Materials and Methods 

This study is based on the mature cypselas from collections in AD, BRI, LISC, 
NSW, RB, SRGH, TAI and Z herbaria as designated in Index Herbariorum 
(Holmgren et al. 1990). Some cypselas were collected by the first author indicated 
as S. Mukherjee from some parts of India also and deposited in the Herbarium 
of the Department of Botany, University of Kalyani, Kalyani - 741235, India 
as indicated by the new herbarium acronym, KAL. A list of all studied taxa and 
their source of origin is given in Appendix I. Studied taxa were classified in tribes 
according to Funk et al. (2009), with nomenclature updated to current knowledge 
(mostly according to the online version of THE INTERNATIONAL PLANT 
NAMES INDEX, Harvard University databases) and the genera and species are 
enumerated in alphabetical sequence. 

For microscopic examination, mature dry cypselas were observed after boiling 
with water and fixed in FAA solution (Johansen 1940). Some FAA preserved 
cypselas were boiled in saturated solution of chloral hydrate solution for 1-3 
minutes and washed with water. Whole cypselas (both treated and untreated) 
were stained in 0.5% aqueous safranin solution and were mounted in 70% phenol 
glycerine solution for microscopic observation. 

Camera lucida drawings were done using a compound trinocular research 
microscope. For each species, at least five randomly selected cypselas were 
examined. 

For scanning electron microscopy (SEM) studies dry cleaned cypselas were placed 
directly on the stubs with double coated adhesive tape and coated with gold. The 



Comp. Newsl. 50, 2012 85 



cypselar surface with or without hairs was scanned and photographed in a Philips 
Electron Microscope at 15 KV in the Regional Sophisticated Instrumentation 
Centre of Bose Institute at Kolkata, West Bengal, India. A few SEM photographs 
were taken in Hitachi SEM at the University Scientific Instrumentation Centre 
(USIC) of the University of Burdwan, Burdwan, West Bengal, India. 

Descriptive terminology for trichomes follows Ramayya (1962 a, b, c), Narayana 
(1979), Payne (1978) and Ciccarelli et al. (2007). However, simple self 
explanatory terms are included to identify the specific type of hair. 

Discussion 

Out of 71 studied species, 53 species (75%) have hairs on the mature cypselar 
surface and 18 species (25%) lack hairs. Of those possessing cypselar hairs, only 
six taxa (11%) possess only glandular hairs (G), 33 species (62%) have non- 
glandular hairs (NG), and remaining 14 species (26%) have both glandular and 
non-glandular hairs. 

As indicated above the classification of Compositae has been recently thoroughly 
revised, but is still in some groups in a state of flux. Bremer (1994) in his cladistic 
classification of Compositae mentioned that in spite of the beginning accumulation 
of modern molecular data, a continued detailed study of morphological characters 
is necessary for the construction of robust phylogenies. This statement is still valid 
today. 

Detailed morpho-anatomical studies of cypselas have been performed at least 
since the time of Schulz-Bipontinus (1844 a, b), and important pioneering work 
by Berkhey (1761). Gaertner (1790) and numerous contributions by Henri 
Casseni (1781 - 1832) must also be acknowledged. In later times, many workers 
(e.g., Briquet 1916, 1930, Blake 1918, Giroux 1930, Kynclova 1970, Grau 
1977, 1980, Saenz 1981, Valez 1981, Pandey et al. 1982, Mukherjee 2000, 
2001 a, b, 2008, Das & Mukherjee 2008) have clearly indicated that diacritical 
features of cypselas could be used for the delimitation, characterization and 
phylogenetic relationships among the taxa of Compositae. Important features of 
cypsela include general morphology, presence or absence of ribs, carpopodium, 
stylopodium, pappus, surface trichomes, phytomelanin layer, calcium oxalate 
crystals, and tissue organization of pericarp and testa. All these characters are 
genetically controlled (Lane 1985, Crawford et al. 2001). These micro-characters 
of cypselas are used for characterization of taxa at the generic and specific level 
and also for improvement of existing system of classification, (cf. Kallersjo 
1985, Mukherjee 2000, 2001 a, b, Mukherjee & Sarkar 2001a,b,c, Mukherjee 
& Nordenstam 2004, 2008, 2010). Therefore, cypselar hairs are part of the set of 



Comp. Newsl. 50, 2012 



microcharacters which all play similar roles along with other features. 

Basic structure, distribution and apical part of hairs are all significant at the specific 
level. Morphological similarity in such details often reflects taxonomic closeness 
as pointed out by Anderberg (1989, 1991 a, b) and EldenAs et al. (1996). An 
example from the present study is provided by Crepis and Hypochaeris which 
have identical types of hairs. Both genera belong to the tribe Cichorieae and they 
are related (Clade 4 in Kilian et al. 2009). 

Hess (1938) has argued that the principal function of 'twin hairs' is absorption of 
water, particularly those which have thickened walls. According to him, this type 
of hairs has sufficient amount of pits which facilitate the absorption of humidity 
from surrounding atmosphere. Therefore plants with such type of cypselar hair 
can thrive in arid and semiarid regions. Myxogenic 'twin hairs' keep the moisture 
around the cypsela as a mucilaginous layer, thus facilitating germination and/or 
perhaps contribute to seed dispersal. Present study indicates that phylogenetically 
widely separated tribes (e.g., Mutisieae and Cichorieae) possess twin type of 
hairs, which are also recorded in literature from many different groups of the 
family. So, this phenomenon may represent a convergent evolution in different 
tribes in response to similar environmental conditions. 

Secretary substances within the glandular hairs are thought to protect plants 
against herbivores and pathogens, and at the same time they might reduce the loss 
of water through cuticular transpiration as well as maintaining the temperature of 
the leaves (Dell & McComb 1975, 1977, Kelsey & Shafizadeh 1980, Werker & 
Fahn 1981, Duke et al. 1994, Tattini et al. 2000). Glandular trichomes of Stevia 
rebaudiana contain sesquiterpene lactones, which are widespread throughout the 
family in various organs. Presence of such compounds, which have a bitter taste 
and may be toxic to grazing animals, should be of importance in the protection of 
plants against herbivores. 

According to Mayekiso et al. (2008), the non-glandular trichomes "appeared to 
originate from the epidermal layer by periclinal division. This process continued 
by periclinal division until several cells which formed the uniseriate trichomes, 
was produced". 

Hairs are directly attached to, or in close contact with the surrounding environment, 
so their role must be largely protection from adverse biological, chemical and 
physical conditions. Absorbing and secretary activity of hairs also influence 
pollination and seed dispersal (cf. above, and Uphof 1962, Werker 2000). 

Presence of 'twin hairs' indicates that the studied subfamilies (Mutisioideae, 
Carduoideae, Pertyoideae, Cichorioideae) are apomorphic in relation to the 
primitive subfamily Barnadesioideae, characterized by its 'barnadesioid hairs' 



Comp.Newsl. 50,2012 87 



(Cabrera 1959, 1977, Bremer 1987, Bremer & Jansen 1992, Kaunas & Stuessy 
1997, Gruenstaeudl et al. 2009). These are generally three-celled trichomes often 
with a long apical cell, a basal isodiametric cell, and an attached epidermal cell 
(two parallel cells). 

Variation of twin hair' structure was indicated already by Hess (1938) and further 
discussed by Freire & Kaunas (1995). Present study also shows the variation of 
twin hairs in different taxa. 

Biseriate (usually 1 0-celled) glandular hairs are common in Compositae (Carlquist 
1 958, 1 959a, b, c, 1 96 1 ), but not observed in mature cypselas. Such hairs have been 
reported from various tribes of several subfamilies, e.g. Mutisieae (Castro et al. 
1997); Vernonieae (Narayana 1979, Castro et al. 1997) and Cardueae (Schnepf 
1969). Peltate glands are very prominent in different species of Vernonia. These 
are visible with simple dissecting microscope. Such glandular hairs were termed 
as 'bilobed hairs' by Faust & Jones (1973), and they are apparently common in 
the tribe Vernonieae. 

Our study also confirmed that mature cypselar hairs of Compositae are often good 
taxonomic markers and can be utilized to resolve the taxonomic affinity of a group. 
For example, the tribe Vernonieae is mainly characterized by the presence of 
both glandular and non-glandular hairs in many studied genera, e.g. Centrapalis, 
Lepidaploa, Polydora, Vanillosmopsis, Vernonanthura and Vernonia. Previously 
almost all these taxa have been included in the genus Vernonia (e.g., Jones 1977). 

Ciccarelli et al. (2007) discussed the distribution of hairs on the ovary surface 
in different taxa of Compositae. The distribution of hairs is perhaps remaining 
constant within a species. Shape of the hair tip is greatly variable and sometimes 
species specific according to the documentation by Ciccarelli et al. (I.e.). Our 
findings agree with these observations. 

Examination with the SEM revealed the presence of two types of hairs, 
corresponding to the two general types recognized in many previous works (cf 
above). The non-secreting or non-glandular type is abundant and responsible 
for covering or protecting the cypselar surface. Secreting hairs are usually club- 
shaped or spheroidal glandular structures of varying size and shape. The cuticular 
striations are either smooth or reticulate in nature. On the basis of this character two 
species of Cicerbita (C. cyanea and C. macrorhiza) can be easily distinguished. 

Narayana (1979) has reported three types of trichomes from the cypselas of 
Vernonia. He designated the 'twin hair' as 'biseriate forked hair'. His other two 
types of hairs have not been observed in the present study. Out of six hair types 
from 20 species of Vernonia reported by Sahu (1984), only the 'twin hair' was 
observed in the present study (by Sahu designated as "achenial hair"). The peltate 



Comp. Newsl. 50, 2012 



gland is usually found mainly on the leaves of Vernonia spp. This type of gland 
was designated as 'bilobed trichome' by Faust & Jones (1973), and also found 
on the mature cypselas of different taxa of Vernonieae. Actually this gland is 
composed of 10 cells (cf above). Such type of trichome has been noted by Favi et 
al. (2008) from Vernonia galamensis ssp. galamensis var. ethiopica. 

In Arctotheca, the entire cypsela is completely embedded in white indumentum 
formed by filiform, entangled hairs with simple obtuse apex but not 'twin' type, 
while in Arctotis, the major portion of cypselar surface is glabrous, except near the 
base, where it possesses numerous, white, 3-5.5 mm long 'twin hairs', whereas 
in Berkheya the entire cypselar surface bears 'twin hairs' of varying size. On 
the basis of these hair characters these three genera of Arctoteae can be easily 
distinguished. 

According to Bremer (1994) and earlier system of classifications (Bentham 
1873a,b, Cabrera 1977) Ainsliaea, Gerbera and Dicoma belonged to the tribe 
Mutisieae. But on the basis of recent studies (Panero & Funk 2002, Kadereit & 
Jeffrey 2007, Funk et al. 2009) these three genera have been included in three 
distinct tribes. Gerbera remains in Mutisieae (subfamily Mutisioideae), whereas 
Dicoma (and Macledium Cass, now recognized as separate) belong in the tribe 
Dicomeae of the subfamily Carduoideae and Ainsliaea in the tribe Pertyeae under 
the subfamily Pertyoideae. If we look into structure of hairs on cypsela, these 
three taxa each have a distinct type of hairs as well as a distinct type of cypselar 
tissue organization (Mukherjee 2001a). These findings support the separation of 
these and related taxa into three different tribes. 

In Cardueae, cypselar surface is often bossed, seldom cross rugose (Arctium), 
covered by unicellular hairs (Centaurea cyanus), possessing scabrous multicellular 
hairs (Echinops) or carrying 'twin hairs' (Carlina) or glabrous in others. Majority 
of aforesaid types except unicellular hairs have been mentioned by Dittrich 
(1977) from this tribe. The unicellular hair on the cypselar surface of Centaurea 
has been reported by Briquet (1930), however. A small swollen and glandular 
hair-like outgrowth is seen at the base of each side of the cypsela in Cirsium 
japonicum. Such type of hair has not been found in other studied species of this 
tribe. Petit (1997) has used cypselar trichomes along with other morphological 
characters to elucidate relationships in Cardueae. 

In mature cypselas, variation of hairs is significantly less than in the young 
cypsela or in the vegetative organs. Only seven basic types of hairs have been 
observed in the present study, whereas numerous and various types of trichomes 
have been noted by different workers either from the vegetative part or from the 
young reproductive parts in other taxa. The basic types of hairs observed by us 
are as follows: 



Comp. Newsl. 50, 2012 



1. Twin hairs; 2. Papillate hairs; 3. Vesicle-like capitate glandular hairs; 
4. Non-glandular filiform hairs; 5. Multicellular non-glandular acroscopic hairs; 
6. Simple hairs; 7. Unicellular hairs. 

Among these the twin hairs, papillate types and vesicle-like capitate glandular hairs 
are predominant. This is perhaps due to their persistent nature as compared to the 
deciduous nature of other types of hairs. Similarly detailed cellular configuration 
of hairs is not clearly visible in mature state of the cypselas. Therefore the detailed 
cellular structure of hairs has not been included here. 

1. Twin type of hairs 

These are prevalent in studied tribes Dicomeae, Pertyeae, Mutisieae, Cichorieae 
and Vernonieae, but less common in the tribes Arctoteae and Cardueae. Twin hairs 
again can be broadly categorized on the basis of shape of the terminal and basal 
cells into the following subtypes: 

i) Long twin hairs : consisting of long laterally adpressed cells: Ainsliaea 
latifolia, A. reflexa, Berkheya zeyheri, Carlina vulgaris, Catananche 
caerulea, Macledium sessiliflorum, Vernonanthura diffusa, Vernonia 
scorpioides, 

ii) Short twin hairs : arm or apical cells are short in length: Gerbera 
jamesonii, Vernonia glabra. 

iii) Apical ends of the arm or apical cells at distinctly unequal levels: Ainsliaea 
latifolia, Baccharoides calvoana ssp. meridionalis, Vernonanthura diffusa, 
Vernonia cistifolia. 

iv) Apical ends of the arm or apical cells more or less in equal plane: 
Catananche caerulea , Elephantopus scaber, Lepidaploa gracilis, 
Polydora bainesii, Polydora poskeana, Vernonia glabra, Vernonia 
scorpioides. 

v) Free portion of arm cells or apical cells greater in length than the basal 
united portion: Baccharoides anthelmintic a, Centrapalis kirkii. 

vi) Papillate twin hairs: two papilla-like cells united to form twin hair: 
Cicerbita cyanea (only found in ribs region). 

vii) Fusiform twin hairs: arm cell thin-walled, more or less equal in length with 
abruptly attenuated tips, usually coloured (pink): Piloselloides hirsuta. 

viii) Base of twin hairs bilobed: Gerbera jamesonii . 

ix) Basal cells of twin hairs distinctly recurved: Macledium sessiliflorum. 

On the basis of orientation or distribution of twin hairs on the cypselar surface, 



90 Comp. Newsl. 50, 2012 



hairs may be following types. 

i) Densely distributed throughout the cypselar surface, e.g., Carlina 

acanthifolia, Vernonia cistifolia, Vernonia glabra, Vemonia scorpioides . 

ii) Sparsely distributed throughout the cypselar surface, e.g., Lepidaploa 
gracilis. 

iii) Restricted (mainly) to furrow region, e.g., Linzia melleri. 

iv) Restricted (mainly) to ribs region, e.g., Polydora bainesii, Vernonia petersii. 

v) Mainly concentrated towards the lower part of cypsela, e.g., Gerbera 
jamesonii. 

vi) Mainly concentrated towards the apical part of cypsela, e.g., Cicerbita 
macrorhiza. 

2. Papillate hairs 

Papillate hairs are usually present in the majority of studied taxa of the tribe 
Cichorieae. On the basis of structure of other types of hairs and distribution of 
papillate hairs on the cypselar surface, the members of Cichorieae can be grouped 
into the following types: 

i) Papillate hairs moderately long with granule-like micro-projections as 
secondary sculpture, e.g., Cicerbita cyanea, Lactuca serriola. 

ii) Papillate hairs without micro-projections, e.g., Actites megalocarpa, 
Cicerbita macrorhiza, Taraxacum officinale. 

iii) Papillate hairs extremely minute and randomly dispersed, e.g., Hieracium 
villosum. 

iv) Numerous minute papillate hairs arranged in several transverse tires and 
forming a transversely muricate surface, e.g., Scorzoneroides autumnalis. 

v) Papillate hairs free at the furrows, but on the ribs papillate hairs laterally 
connate with each other and forming multicellular scale-like structures. 
Based on the shape of these scale-like structures there are three subtypes: 

a) Apex of the scale nearly flat, e.g., Crepis vesicaria. 

b) Apex of the scale nearly triangular, e.g., Hypochaeris glabra. 

c) Apex of the scale nearly rounded, e.g., Hypochaeris radicata. 

vi) Papillate hairs with wide bifurcate base and pointed apex, e.g., Taraxacum 
officinale. 



Comp.Newsl. 50,2012 91 



vii) True papillate hairs absent, but surface squamosely muricate. Muricae 
vertically elongated with rounded broad elevation. Epidermal cells form 
obscurelypapilla-likestructureswithobtuseapex,e.g., Tragopogonporrifolius . 

viii) Cypselar surface with papillate hairs at the furrows and moderately long 
papillate twin hairs on the ribs, e.g., Actites megalocarpa, Cicerbita 
cyanea. 

ix) Cypselar surface without papillate hairs but with long twin hairs, e.g., 
Catananche caernlea. 

x) Cypselar surface without any type of hairs but cross rugose or cross 
marked, e.g., Sonchus brachyotus. 

3. Vesicle-like capitate glandular hairs 

These are prevalent in the tribe Vernonieae. On the basis of distribution and 
structure of these hairs on the cypselar surface the following categories are 
recognized: 

i) Glandular hairs uniformly distributed throughout the cypselar surface, e.g., 
Vernonia colorata. 

ii) Glandular hairs sparsely distributed on the furrows only, and cypselar hairs 
may be of two types: 

a) Glandular hairs homomorphic, e.g., Centrapalis kirkii, Gymnanthemum 
amygdalinum, Lepidaploa gracilis, Polydora bainesii, Vanillosmopsis 
capitata, Vernonanthura diffusa, Vernonia cistifolia, Vernonia petersii, 
Vernonia scorpioides. In Polydora bainesii each glandular hair has three 
distinct zones and in Polydora poskeana each glandular hair has two semi- 
lunar hyaline spaces at the two opposite sides (visible after clearing). 

b) Glandular hairs dimorphic, e.g., Rolandra fruticosa, Vernonia colorata. 

iii) Glandular hairs densely distributed on the furrows of cypsela. There are 
three subcategories: 

a) Glandular hairs arranged in 4-9 vertical zones, e.g., Bothriocline laxa, 
Centrapalis kirkii, Polydora bainesii, Vernonia cistifolia. 

b) Glandular hairs arranged in 1-3 vertical rows, e.g., Gymnanthemum 
amygdalinum, Lepidaploa gracilis, Vanillosmopsis capitata, Vernonanthura 
diffusa. 

c) Glandular hairs arranged in 1-2 vertical rows in two sides of each furrow, 
e.g., Baccharoides calvoana, Linzia melleri, Vernonia glabra. 



92 Comp. Newsl. 50, 2012 



4. Multicellular non-glandular acroscopic hairs. This type is found in only 
one species, viz., Echinops sphaewcephalus. 

5. Unicellular hairs. This unbranched non-glandular hair type was observed 
in Centaurea cyanus and Cirsium vulgare. 

6. Simple hairs, i.e. unbranched multicellular non-glandular hair type, e.g., 
Arctotheca calendula, Arctotis venusta, Centrapalis kirkii, Lactuca serriola, 
Tragopogon porrifolius. 

7. Non-glandular filiform hairs. These hairs have a basal bulbous portion and 
a long filiform terminal part, e.g., Linzia melleri. 

Conclusion 

From the above discussion it is obvious that micro-morphological characters of 
hairs along with other microcharacters of cypsela or other vegetative or floral 
morphological characters can be used for taxonomic and phylogenetic studies 
in the Compositae. There is a need to develop better universal terminology and 
perform detailed comparative studies of these micro-morphological features to 
improve Compositae taxonomy. An intensive morphological and ultrastructural 
study of cypsela hairs of cypsela along with other features of cypsela will make 
it possible to identify species in the fruiting state even when plants in flower are 
not available. 

The present study also suggests that the value of hairs as a taxonomic criterion 
will be greatly increased when combined with other lines of evidence. 



Comp. Newsl. 50, 2012 



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103 



Appendix 1. Taxa studied and specimen location. 



SI. 

No. 


Name of the Species 


Tribe 


Source of Origin 


1. 


Actites megalocarpa 
(HooK.f.) Lander (syn. 
Sonchus megalocarpus 
(HooK.f.) J. M. Black) 


Cichorieae 


AD;A.A. Munir5512 


2. 


Ainsliaea aptera DC. 


Pertyeae 


KAL ; S. Mukherjee 25 


3. 


Ainsliaea latifolia (D. Don) 
Sch.Bip. 


Pertyeae 


KAL ; S. Mukherjee 17 


4. 


Ainsliaea reflexa Merr. var. 
nimbonim Hand.-Mazz. 


Pertyeae 


TAI; Yuh Fong Chen 3300 


5. 


Arctium lappa L. 


Cardueae 


Z;Nr. 343 


6. 


Arctotheca calendula (L.) 
Levyns 


Arctoteae 


AD;N. N. Donner8541 


7. 


Arctotis venusta T. Norl. 


Arctoteae 


Z;Nr. 345 


8. 


Baccharoides anthelmintica 

(L.) MOENCH 


Vernonieae 


KAL; S. Mukherjee 1 


9. 


Baccharoides calvoana 
(HooK.f.) Isawumi, El- 
Ghazaly & B. Nord. 
ssp. meridionalis (Wild) 
Isawumi, El-Ghazaly & B. 
Nord. 


Vernonieae 


SRGH; G.Pope 1930 


10. 


Berkheya zeyheri (Sond. & 
Harv.) Oliv. & Hiern ssp. 
zeyheri 


Arctoteae 


LISC; A.R. Torre 6907 


11. 


Bothriocline laxa N. E. Br. 
ssp. laxa 


Vernonieae 


SRGH; M. Mavi 1 1 


12. 


Carduus defloratus L. 


Cardueae 


Z; Nr. 359 


13. 


Carlina acanthifolia All. 
ssp. cynara (Pourret ex 
Duby) Rouy 


Cardueae 


Z; Nr. 360 


14. 


Carlina vulgaris L. ssp. 
vulgaris 


Cardueae 


Z;Nr. 361 


15. 


Catananche caerulea L. 


Cichorieae 


Z: Nr. 363 


16. 


Centaurea cyanus L. (syn. 
Cyanus segetum Hill) 


Cardueae 


Z;Nr. 364 


17. 


Centaurea macrocephala 
Muss. Puschk. ex Willd. 


Cardueae 


Z;Nr. 366 



104 



Comp. Newsl. 50, 2012 



18. 


Centaurea maculosa Lam. 
ssp. maculosa 


Cardueae 


Z; Nr. 369 


19. 


Centrapalis kirkii (Oliv. & 
Hiern) H. Rob. 


Vernonieae 


LISC; F.A. Mendonca 2033 


20. 


Cicerbita cyanea (D. Don) 
P. Beauv. 


Cichorieae 


KAL; S. Mukherjee 18 


21. 


Cicerbita macrorhiza 
(Royle) R Beauv. 


Cichorieae 


KAL; S. Mukherjee 19 


22. 


Cirsium arvense (L.) Scop. 


Cardueae 


KAL; S. Mukherjee 23 


23. 


Cirsium falconeri (HooK.f.) 
Petr. 


Cardueae 


KAL; S. Mukherjee 12 


24. 


Cirsium japonicum DC. 


Cardueae 


Z; Nr. 378 


25. 


Cirsium vulgare (Savi) Ten. 


Cardueae 


BRI; s.n., s.coll. 


26. 


Crepis pyrenaica (L.) 

Greuter 


Cichorieae 


Z;Nr. 383 


27. 


Crepis vesicaria L. 


Cichorieae 


AD; N.N. Donner 8607 


28. 


Echinops sphaerocephalus 
L. 


Cardueae 


Z;Nr. 386 


29. 


Elephantopus scaber L. 


Vernonieae 


RB; SN 257 


30. 


Gerbera jamesonii Bolus ex 
HooK.f. 


Mutisieae 


Z, Nr. 397 


31. 


Gymnanthemum 
amygdalinum (Delile) 
Sch.Bip. ex Walp. (syn. 
Vernonanthura condensata 
(Baker) H. Rob.) 


Vernonieae 


RB ; SN 249 


32. 


Hieracium villosum Jacq. 


Cichorieae 


Z; Nr. 404 


33. 


Hypochaeris glabra L. 


Cichorieae 


AD; A. A. Munir8601 


34. 


Hypochaeris radicata L. 


Cichorieae 


BRI; s.n., s.coll. 


35. 


Lactuca dissect a D. Don 


Cichorieae 


KAL; S. Mukherjee 43 


36. 


Lactuca graciliflora DC. 
(syn. Stenoseris graciliflora 
(Wall, ex DC.) C. Shih) 


Cichorieae 


KAL; S. Mukherjee 52 


37. 


Lactuca serriola L. 


Cichorieae 


BRI; s.n., s.coll. 


38. 


Launaea acaulis (Roxb.) 
Babc. ex Kerr 


Cichorieae 


KAL; S. Mukherjee 39 


39. 


Launaea aspleniifolia 
(Willd.) HooK.f. 


Cichorieae 


KAL; S. Mukherjee 54 



Comp. Newsl. 50, 2012 



105 



40. 


Launaea procumbens 
(Roxb.) Ramayya & 
Rajagopal 


Cichorieae 


KAL; S. Mukherjee 46 


41. 


Launaea sarmentosa 
(Willd.) Sch.Bip. ex Kuntze 


Cichorieae 


KAL; S. Mukherjee 37 


42. 


Leibnitzia nepalensis 
(Kunze) Kjtam. 


Mutisieae 


KAL ; S. Mukherjee 44 


43. 


Lepidaploa gracilis (Kunth) 
H. Rob. 


Vernonieae 


RB; SN 250 


44. 


Linzia melleri (Oliv. & 
Hiern) H. Rob. 


Vernonieae 


LISC; R.Santos 2051 


45. 


Macledium sessiliflorum 
(Harv. in Harv. & Sond.) S. 
Ortiz ssp. sessiliflorum 


Dicomeae 


LISC;A.R. Torre 13 


46. 


Picris hieracioides L. 


Cichorieae 


KAL; S. Mukherjee 29 


47. 


Piloselloides hirsuta 
(Forssk.) C. Jeffrey ex 
Cufod. 


Mutisieae 


KAL; S. Mukherjee 28 


48. 


Polydora bainesii (Oliv. & 
Hiern) H. Rob. 


Vernonieae 


SRGH; G.Pope 1929 


49. 


Polydora poskeana (Vatke 
& Hildebrandt) H. Rob. 


Vernonieae 


LISC; A. R. Torre & Paiva 
11332 


50. 


Prenanthes khasiana C. B. 
Clarke 


Cichorieae 


KAL ; S. Mukherjee 67 


51. 


Pseudelephantopus spicatus 
(Juss.) C. F. Baker 


Vernonieae 


KAL; S. Mukherjee 21 


52. 


Ptilostemon diacantha 
(Labill.) Greuter 


Cardueae 


Z; Nr. 377 


53. 


Rhaponticum scariosum 
Lam. ssp. rhaponticum 
(L.) Greuter (syn. Leuzea 
rhapontica (L.) J. Holub) 


Cardueae 


Z;Nr. 413 


54. 


Rolandra fruticosa (L.) 
Kuntze 


Vernonieae 


RB; SN 255 


55. 


Saussurea abnormis Lipsch. 
(syn. Himalaiella abnormis 
(Lipsch.) Raab-Straube, 
should prob. be treated in 
Jurinea) 


Cardueae 


KAL; S. Mukherjee 32 


56. 


Saussurea deltoidea (DC.) 
Sch. Bip. 


Cardueae 


KAL; S. Mukherjee 22 



106 



Comp.Newsl. 50,2012 



57. 


Sanssiirea heteromalla (D. 
Don) Hand.-Mazz. 


Cardueae 


KAL; S. Mukherjee 27 


58. 


Scorzoneroides autumnalis 
(L.) Moench (syn. 
Leontodon autumnalis L.) 


Cichorieae 


Z; Nr. 409 


59. 


Sonchus brachyotus DC. 


Cichorieae 


KAL; S. Mukherjee 20 


60. 


Sonchus schweinfurthii 
Oliv. & HlERN 


Cichorieae 


SRGH; M. Mavi 8 


61. 


Taraxacum officinale Weber 


Cichorieae 


AD; A. A. Munir 5500 


62. 


Tarlmounia elliptica (DC.) 
H. Rob., S. C. Keeley, 
Skvarla & R. Chan (syn. 
Vernonia elliptica DC.) 


Vernonieae 


KAL; S. Mukherjee 57 


63. 


Tragopogon porrifolius L. 


Cichorieae 


AD; N. N. Donner 8606 


64. 


Vanillosmopsis capitata 
(Spreng.) Sch. Bip. 


Vernonieae 


RB; SN 248 


65. 


Vernonanthura diffusa 
(Less.) H. Rob. 


Vernonieae 


RB; SN 254 


66. 


Vernonia cinerea (L.) Less. 


Vernonieae 


KAL; S. Mukherjee 55 


67. 


Vernonia cistifolia O. Hoffm. 


Vernonieae 


SRGH; G.Pope 1931 


68. 


Vernonia colorata (Willd.) 
Drake ssp. colorata (syn. V. 
senegalensis Less.) 


Vernonieae 


LISC; Schlieben 2457 


69. 


Vernonia glabra (Steetz) 
Vatke 


Vernonieae 


SRGH ; M. Mavi 12 


70. 


Vernonia peter sii Oliv. & 
Hiern ex Oliv. 


Vernonieae 


LISC; A. R.Torre 118 


71. 


Vernonia scorpioides (Lam.) 
Pers. 


Vernonieae 


RB;SN251 



Comp. Newsl. 50, 2012 



107 




Figs. l-38.The structure and distribution of hairs from mature cypselas in 
different taxa. 



108 Comp. Newsl. 50, 2012 



Figs. 1,2: Gerbera jamesonii. 

Fig. 3: Piloselloides hirsuta. 

Figs. 4, 5, 6: Macledium sessiliflorum ssp. sessilifloram. 

Fig. 7: Arctium lappa. 

Figs. 8, 9: Carlina acanthifolia ssp. cynara. 

Figs. 10, 11: Carlina vulgaris ssp. vulgaris. 

Figs. 12, 13: Centaurea cyanus. 

Fig. 14: Cirsium japonicum . 

Fig. 15: Cirsium vulgare. 

Figs. 16, \1 : Echinops sphaerocephalus . 

Fig. 18: Saussurea deltoidea. 

Figs. 1-9, 20: Ainsliaea latifolia. 

Fig. 21: Actites megalocarpa. 

Fig. 22: Catananche caerulea. 

Fig. 23: Cicerbita cyanea. 

Figs. 24-28: Cicerbita macrorhiza. 

Fig. 29: Crepis vesicaria. 

Fig. 30: Hieracium villosum. 

Fig. 3 1 : Hypochaeris glabra. 

Figs. 32, 33: Hypochaeris radicata. 

Fig. 34: Lactuca serriola. 

Fig. 35: Scorzoneroides autumnalis. 

Fig. 36: Taraxacum officinale. 

Fig. 37: Tragopogon porrifolius . 

Fig. 38: Arctotheca calendula. 



Comp.NewsI. 50,2012 



109 




Figs. 39-84. The structure and distribution of hairs from mature cypselas 
in different taxa. 



110 Comp. Newsl. 50, 2012 



Fig. 39: Arctotis venusta. 

Fig. 40: Berkheya zeyheri ssp. zeyheri. 

Fig. 41: Baccharoides anthelmintica. 

Figs. 42, 43: Baccharoides calvoana ssp. meridionalis . 

Figs. 44, 45: Bothriocline laxa ssp. /ara. 

Figs. 46^8: Centrapalis kirkii. 

Fig. 49: Elephantopus scaber. 

Figs. 50—52: Lepidaploa gracilis . 

Figs. 53-55: Linzia melleri. 

Figs. 56-58: Polydora bainesii. 

Figs. 59-61: Polydora poskeana. 

Figs. 62, 63: Rolandra fruticosa. 

Figs. 64, 65: Vanillosmopsis capitata. 

Figs. 66, 67: Gymnanthemum amygdalinum. 

Figs. 68-70: Vernonanihura diffusa. 

Figs. 71-73: Vernonia cistifolia. 

Figs. 74-76: Vernonia glabra. 

Figs. 77, 78: Vernonia petersii . 

Figs. 79-81: Vernonia s cor pioides. 

Figs. 82-84: Vernonia colorata ssp. colorata. 



Comp. Newsl. 50, 2012 



111 




Figs. 85-96. SEM photographs showing the structure and distribution 
of cypselar hairs. 

Fig. 85: Actites megalocarpa, x 400. Fig. 86: Ainsliaea latifolia, x 200. 

Fig. 87: Ainsliaea reflexa, x 100. Fig. 88: Arctium lappa, x 50. 

Fig. 89: Arctotheca calendula, x 50. Fig. 90: Arctotis venusta, x 50. 

Fig. 91: Baccharoides anthelmintica. 

Fig. 92: Baccharoides calvoana ssp. meridionalis, x 1600. 

Fig. 93: Berkheya zeyheri, x 50. 

Figs. 94, 95: Bothriocline laxa, x 100; x 1600. 

Fig. 96: Carlina acanthifolia, x 100. 



112 



Comp. Newsl. 50, 2012 




Figs. 97-108. SEM photographs showing the structure and distribution of 
cypselar hairs. 

Fig. 97: Centaurea cyanus, x 25. Fig. 98: Cicerbita cyanea, x 800. 

Fig. 99: Cicerbita macrorhiza, x 800. Fig. 100: Crepis vesicaria, x 400. 

Fig. 101: Echinops sphaerocephalus, x 400. Fig. 102: Hieracium villosum, x 

800. Fig. 103: Hypochaeris glabra, x 400. Fig. 104: Hypochaeris radicata, x 

400. 

Fig. 105: Lactuca serriola, x 1600. Fig. 106: Scorzoneroides autumnalis, x 400. 

Fig. 107: Linzia melleri, x 400. Fig. 108: Macledium sessiliflorum ssp. 

sessiliflorum, x 400. 



Comp. Newsl. 50, 2012 



113 




Figs. 109-120. SEM photographs showing the structure and distribution of 
cypselar hairs. 

Figs. 109, 110: Polydora bainesii, x 50; x 400. Figs.lll, 112: Rolandra 

fruticosa. 

Fig. 113: Sonchus brachyotus, x 400. Fig. 114: Tragopogon porrifolius, x 400. 

Fig. 115: Vanillosmopsis capitata, x 50. Fig. 116: Vernonia cistifolia, x 800. 

Fig. 117: Vernonia glabra, x 200. Fig. 118: Vernonia peter sii, x 400. 

Fig. 119: Vernonia scorpioides, x 100. Fig. 120: Vernonia colorata ssp. 

colorata, x 400. 



114 Comp. Newsl. 50, 2012 



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New taxa and combinations published 
in this issue 

Caputia B. Nord. & Pelser, gen. nov.: p. 59 

Caputia medley-woodii (Hutch.) B. Nord. & Pelser, comb, nov.: p. 62 

Caputia pyramidata (DC.) B. Nord. & Pelser, comb, nov.: p. 67 

Caputia scaposa (DC.) B. Nord. & Pelser, comb, nov.: p. 66 

Caputia scaposa var. addoensis (Compton) B. Nord. & Pelser, comb, nov.: 
p. 67 

Caputia scaposa var. caulescens (Harv.) B. Nord. & Pelser, comb, nov.: p. 67 

Caputia tomentosa (Haw.) B. Nord. & Pelser, comb, nov.: p. 65 

Crassothonna B. Nord., gen. nov.: p. 71 

Crassothonna alba (Compton) B. Nord., comb, nov.: p. 72 

Crassothonna cacalioides (L.f.) B. Nord., comb, nov.: p. 72 

Crassothonna capensis (L. H. Bailey) B. Nord., comb, nov.: p. 72 

Crassothonna clavifolia (Marl.) B. Nord., comb, nov.: p. 73 

Crassothonna cylindrica (Lam.) B. Nord., comb, nov.: p. 73 

Crassothonna discoidea (Oliv. in Hook.) B. Nord., comb, et stat. nov.: p. 73 

Crassothonna floribunda (Schltr) B. Nord., comb, nov.: p. 73 

Crassothonna opima (Merxm.) B. Nord., comb, nov.: p. 74 

Crassothonna patula (Schltr) B. Nord., comb, nov.: p. 74 

Crassothonna protecta (Dinter) B. Nord., comb, nov.: p. 74 

Crassothonna rechingeri (B. Nord.) B. Nord., comb, nov.: p. 75 

Crassothonna sedifolia (DC.) B. Nord., comb, nov.: p. 75 

Crassothonna sparsiflora (S. Moore) B. Nord., comb, nov.: p. 75 



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